Skip to main content
PMC home page
Biodiversity Data Journal logoLink to Biodiversity Data Journal
. 2021 Dec 22;9:e75907. doi: 10.3897/BDJ.9.e75907

Tylocinum is no longer monotypic: Tylocinumbrevisporum sp. nov. (Boletales, Boletaceae) from northern Thailand

Bhavesh Raghoonundon 1,2, Naveed Davoodian 3, Monthien Phonemany 1,2, Olivier Raspé 2,✉
PMCID: PMC8716511  PMID: 34975279

Abstract

Background

Tylocinum Y.C. Li & Zhu L. Yang 2016 is a Boletaceae genus belonging in subfamily Leccinoideae. It was described in 2016 from China and, prior to this study, it contained only one species, T.griseolum Y.C. Li & Zhu L. Yang 2016. During our survey of Boletaceae from Thailand, we collected some specimens that could be identified as a Tylocinum species, different from T.griseolum.

New information

The bolete specimens, collected in forests dominated by Dipterocarpaceae and Fagaceae in northern Thailand, are described as Tylocinumbrevisporum Raghoonundon & Raspé sp. nov. Macroscopic and microscopic descriptions with illustrations are provided, as well as a 3-gene phylogeny, which confirms the new taxon’s position in Tylocinum. Tylocinumbrevisporum differs from the only other known Tylocinum species (T.griseolum) by its brownish-grey colour, greyish-orange to brownish-orange colour change in the hymenophore when bruised, smaller pores (≤ 0.5 mm), longer tubes (up to 6 mm long), shorter and narrower basidiospores, longer and broader basidia and longer pleurocystidia relative to cheilocystidia. T.brevisporum is the second species from the genus Tylocinum and the only one to be found outside China thus far.

Keywords: new species, Boletaceae, Leccinoideae, molecular phylogeny, taxonomy, Thailand

Introduction

Tylocinum Y.C. Li & Zhu L. Yang 2016, is a monotypic genus of ectomycorrhizal (ECM) boletes (Boletaceae, Boletales, Agaricomycetes, Basidiomycota, Fungi). Typical characters of the genus are its dark scabrous stipe surface, white to pallid unchanging context in the pileus and stipe, white to pallid hymenophore, trichodermium pileipellis and smooth basidiospores (Wu et al. 2016). The type species Tylocinumgriseolum Y.C. Li & Zhu L. Yang 2016, was originally described from China and was the only species known from this genus at the time. The phylogenetic analyses by Wu et al. (2016) showed that Tylocinum forms a separate clade from all other generic clades in the subfamily Leccinoideae.

The plant family Dipterocarpaceae includes many species of large trees that are often dominant in the tropical and subtropical lowlands of Southeast Asia, where the species diversity of Dipterocarpaceae is highest (Ashton 1982, Hamilton et al. 2019). Many Dipterocarpaceae are well known to be ECM, symbiotically associating with various ECM fungi, including mushroom-forming species (Watling et al. 2002, Yuwa-Amornpitak et al. 2006, Brearley 2012). Several new genera and species of boletes have recently been documented from tropical dipterocarp forest (Desjardin et al. 2009, Neves et al. 2012, Hosen et al. 2013, Halling et al. 2014, Raspé et al. 2016, Wu et al. 2016, Vadthanarat et al. 2019, Chuankid et al. 2019). Members of the Fagaceae, which also form ECM associations, co-occur with dipterocarps in Southeast Asia (Smith et al. 2008), which promotes higher mycodiversity and ECM colonisation in those tropical forest ecosystems (Corrales et al. 2018).

In this study, we describe a new species of Tylocinum from dry dipterocarp forests of northern Thailand, with description, illustrations and molecular phylogenetic analyses of a multi-gene DNA sequence dataset (atp6, tef1 and rpb2).

Materials and methods

Specimens collected

Fresh basidiomata were collected during the rainy season (2019) from Chiang Mai and Chiang Rai Provinces, orthern Thailand. The basidiomata were photographed on-site and wrapped in aluminium foil. The descriptions of the macroscopic features were made on the same day, after which the basidiomata were dried in an electric drier at 45–50°C. Specimens were deposited in the Mae Fah Luang University (MFLU) or CMUB Herbaria.

Ecological, morphological and taxonomic study

The habitat, locality information and macro-chemical reactions on fresh basidiomata were recorded. Spore prints were taken for each collection. Colour codes were given using Kornerup and Wanscher (1978) as a guide. Microscopic characters were studied in the dried specimens. The following mounting solutions were used to observe the tissues: 10% aqueous potassium hydroxide (KOH) or 28–30% ammonium hydroxide (NH4OH) solutions or 1% ammoniacal Congo red solution. The microscopic structures were studied at magnifications of 60× and 100×, photographed with a calibrated Nikon Y-TV55 camera, fitted to a Nikon DIC microscope. A total of 60 basidiospores, 30 basidia, 30 pleurocystidia, 30 cheilocystidia and 30 terminal cells and 30 hyphae for both the pileipellis and stipitipellis were measured. The dimensions of the microscopic features 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 for the spores, is presented in the same format. All microscopic features were drawn by free hand, using a drawing tube. Faces of Fungi (Jayasiri et al. 2015) and MycoBank numbers are provided for the new species.

DNA extraction, PCR amplification and sequencing

Genomic DNA was extracted from CTAB-preserved tissues or dry specimens (ca. 10 mg) using a CTAB isolation procedure, adapted from Doyle and Doyle (1990). The atp6, tef1 and rpb2 gene regions were amplified by polymerase chain reaction (PCR). For amplification of atp6, the primers ATP6-1M40F and ATP6-2M were used (Raspé et al. 2016). EF1-983F and EF1-2218R (Rehner and Buckley 2005) were used to amplify tef1 and bRPB2-6F and bRPB2-7.1R (Matheny 2005) were used to amplify rpb2. The PCR amplification, purification and sequencing of atp6, rpb2 and tef1 were used following the procedure from Raspé et al. (2016).

Sequence alignment and phylogenetic analysis

The sequences were assembled using Geneious 8 (Biomatters). The Basic Local Alignment Search Tool (BLAST) (https://blast.ncbi.nlm.nih.gov/Blast.cgi) from GenBank was used to find the closest matches to the sequences. Reference sequences (Table 1) were downloaded and aligned using MAFFT v. 7 (Katoh and Standley 2013; http://mafft.cbrc.jp/alignment/server/). Then, the concatenated three-gene matrix was prepared.

Table 1.

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

Species Voucher Origin atp 6 tef 1 rpb 2 References
Baorangiamajor OR0209 Thailand MG897421 MG897431 MG897441 Phookamsak et al. (2019)
Baorangiapseudocalopus HKAS75739 China – KJ184570 KM605179 Wu et al. (2015)
Baorangiarufomaculata BOTH4144 USA MG897415 MG897425 MG897435 Phookamsak et al. (2019)
Borofutusdhakanus OR0345 Thailand MH614660 MH614709 MH614755 Vadthanarat et al. (2018)
Ionosporuslongipes LEE1180 Malaysia MT085461 MT085471 MH712031 Khmelnitsky et al. (2019)
Lanmaoaasiatica OR0228 China MH614682 MH614730 MH614777 Vadthanarat et al. (2019)
Lanmaoacarminipes BOTH4591 USA MG897419 MG897429 MG897439 Phookamsak et al. (2019)
Lanmaoapallidorosea BOTH4432 USA MG897417 MG897427 MG897437 Phookamsak et al. (2019)
Leccinummonticola HKAS76669 China – KF112249 KF112723 Wu et al. (2014)
Leccinumquercinum HKAS63502 China – KF112250 KF112724 Wu et al. (2014)
Leccinumscabrum RW105a Belgium KT823979 KT824045 KT824012 Raspé et al. (2016)
Leccinumscabrum VDKO0938 Belgium MG212549 MG212593 MG212635 Vadthanarat et al. (2018)
Leccinumschistophilum VDKO1128 Belgium KT823989 KT824055 KT824022 Raspé et al. (2016)
Leccinumvariicolor HKAS57758 China – KF112251 KF112725 Wu et al. (2014)
Leccinumvariicolor VDKO0844 Belgium MG212550 MG212594 MG212636 Vadthanarat et al. (2018)
Leccinellumaff.crocipodium HKAS76658 China – KF112252 KF112728 Wu et al. (2014)
Lecinellumcf.intusrubens OR0082 Thailand MZ803019 MZ803024 MZ824749 This study
Leccinellumcrocipodium VDKO1006 Belgium KT823988 KT824054 KT824021 Raspé et al. (2016)
Leccinellumcremeum HKAS90639 China – KT990781 KT990420 Wu et al. (2016)
Leccinellum sp. HKAS53427 China – KF112253 KF112727 Wu et al. (2014)
Leccinellum sp. OR0711 Thailand MH614685 MH614733 MH614780 Vadthanarat et al. (2019)
Octavianiahesperi KPM-NC 17793 Japan KC552150 JN378422 – Orihara et al. (2016)
Octavianiajaponimontana KPM-NC 17797 Japan KC552151 JN378425 – Orihara et al. (2016)
Octavianianonae KPM-NC 17748 Japan KC552143 JN378403 – Orihara et al. (2016)
Octavianiatasmanica MEL 2341996 Australia KC552156 JN378436 – Orihara et al. (2012), Orihara et al. (2016)
Octavianiazelleri MES270 USA KC552161 JN378440 – Orihara et al. (2012), Orihara et al. (2016)
Pseudoaustroboletuscf.valens OR0477 China MZ803020 MZ803025 MZ824750 This study
Retiboletusbrevibasidiatus OR0570 Thailand MT085469 MT085476 MT085479 Chuankid et al. (2021)
Retiboletusbrunneolus HKAS 52680 China – KF112179 KF112690 Wu et al. (2014)
Retiboletusfuscus OR0231 China MG212556 MG212600 MG212642 Vadthanarat et al. (2018)
Retiboletusfuscus OR0738 Thailand MT085462 MT085472 MT085477 Chuankid et al. (2021)
Retiboletusgriseus MB03-079 USA KT823964 KT824030 KT823997 Raspé et al. (2016)
Retiboletuskauffmanii OR0278 China MG212557 MG212601 MG212643 Vadthanarat et al. (2018)
Retiboletusnigrogriseus BC0179 Thailand MT085464 MT085474 MT085478 Chuankid et al. (2021)
Retiboletusnigrogriseus OR049 Thailand KT823967 KT824000 KT824033 Raspé et al. (2016)
Retiboletusornatipes MBsn USA MT219514 MT219516 MT219515 Chuankid et al. (2021)
Rhodactinarostratispora SV170 Thailand MG212560 MG212605 MG212645 Vadthanarat et al. (2018)
Rossbeeveraeucyanea TUMH-40252 Japan KC552116 KC552069 – Orihara et al. (2016)
Rossbeeveragriseovelutina TUMH-40266 Japan KC552121 KC552073 – Orihara et al. (2016)
Rossbeeveravittatispora A.W. Claridge 2137 Australia KC552105 KC552063 – Orihara et al. (2016)
Spongiformathailandica DED7873 Thailand MG212563 KF030436 MG212648 Nuhn et al. (2013),
Vadthanarat et al. (2018)
Spongisporatemasekensis ACMF5 Singapore MZ803018 MZ803023 MZ824748 This study
Turmalineamesomorphasubsp.mesomorpha KPM-NC 18012 Japan KC552139 KC552090 – Orihara et al. (2016)
Turmalineapersicina KPM-NC 18001 Japan KC552130 KC552082 – Orihara et al. (2016)
Turmalinea sp. Muroi361 USA DQ218885 DQ219224 DQ219046 Orihara et al. (2016)
Tylocinumgriseolum HKAS50281 China – KF112284 KF112730 Wu et al. (2014)
Tylocinumbrevisporum OR622 Thailand MZ803021 – MZ824751 This study
Open in a new tab

All analyses were done on the CIPRES Science Gateway (https://www.phylo.org; Miller et al. 2012). Maximum Likelihood (ML) phylogenetic tree inference was done using RAxML-HPC2 v.8.2.10 (Stamatakis 2006), using the GTRCAT model of sequence evolution with 25 categories. Three Lanmaoa species and three Baorangia species were selected as outgroup. Four partitions were defined: atp6, tef1 exons, rpb2 exons and introns. Statistical support of the clades was obtained using 1,000 rapid bootstrap replicates.

Using jModeltest2 (Darriba et al. 2012) on XSEDE via the CIPRES Science Gateway, the best-fit model of substitution for analysis in MrBayes was estimated for each gene, based on the Bayesian Information Criterion (BIC). GTR + I + G for atp6 and introns, SYM + I + G for tef1 exons and K80 + I + G for rpb2 exons were selected as the best fit models. Partitioned Bayesian analysis was performed with MrBayes 3.2.7a (Ronquist et al. 2012). Two runs of four cold and one heated chains were run for 1,000,000 generations and sampled every 200 generations. The average standard deviation of split frequencies was 0.005106 at the end of the runs. The burn-in phase (25%) was estimated by checking the stationarity in the plot generated by the sump command.

Taxon treatments

Tylocinum brevisporum

Raghoonundon & Raspé sp. nov.

64D99B61-7573-598D-8BE9-02D0AC73258B

MB841102

FoF 10255

Materials

  1. Type status: Holotype. Taxon: kingdom: Fungi; phylum: Basidiomycota; class: Agaricomycetes; order: Boletales; family: Boletaceae; genus: Tylocinum; specificEpithet: brevisporum; taxonRank: species; Location: country: Thailand; stateProvince: Chiang Rai Province, Chang Wat, Doi Pui; verbatimElevation: 730 m; verbatimCoordinates: 19°48'50"N, 99°51'57"E; Identification: identifiedBy: Bhavesh Raghoonundon; Event: eventDate: 20 August 2019; Record Level: institutionID: MFLU 21-0144; institutionCode: Mae Fah Luang University Herbarium; collectionCode: BR137

  2. Type status: Other material. Taxon: kingdom: Fungi; phylum: Basidiomycota; class: Agaricomycetes; order: Boletales; family: Boletaceae; genus: Tylocinum; specificEpithet: brevisporum; taxonRank: species; Location: country: Thailand; stateProvince: Chiang Mai Province, Mueang District; verbatimElevation: 450 m; verbatimCoordinates: 18°48'40"N, 98°56'31"E; Identification: identifiedBy: Olivier Raspé; Event: eventDate: 18 May 2015; Record Level: institutionID: CMU-B OR622; collectionID: OR622; institutionCode: Chiang Mai University Herbaria

Description

Basidiomata pileo-stipitate, small to medium-sized (Fig. 1). Pileus (1.5–)2.0–2.5 cm in diameter, convex when young, becoming plano-depressed with age; margin deflexed to uplifted, surface finely tomentose, dull and dry, at first brown (7E4) to greyish-brown (8E3–8F4), becoming paler (8D3) near the margin with age; context 3–5 mm thick halfway to the margin, soft and fleshy, off-white, slightly browning on exposure. Stipe central, cylindrical, (3.4–)4.9–6.5 cm × 0.6–1.3 cm, surface even, dull and dry, scabrous, covered with granular squamules (dotted-verrucose), brownish-grey (7E2–8E2) when young to reddish-brown (8E5) to dark brown (8F5) with age, no colour change when bruised, basal mycelium off-white; context solid, fleshy, off-white, reddish-brown to dark brown near the stipe base (8F7) and in worm wounds, slightly browning on exposure. Hymenophore tubulate, subventricose, adnexed, slightly depressed around apex of the stipe, greyish-orange to brownish-orange when bruised. Tubes 3–6 mm long halfway to the margin, off-white, easily separable from one another. Pores ≤ 0.5 mm wide at mid-radius, regularly arranged, angular, off-white, turning brown to dark brown (8E5–8F5) when bruised. Odour fungoid. Taste bitter. Spore print not obtained.

Figure 1.

Figure 1.

Open in a new tab

Photograph of Tylocinumbrevisporum sp. nov. a, b Basidioma of specimen OR622; c Basidioma of the holotype (BR 137).

Basidiospores (6.7–)7.5–10–11.7(–11.8) × (3.1–)3.5–4.7–5.8(–5.9) µm (n = 50) Q = (1.7–) 1.79–2.15–2.5 (–2.61), ellipsoid in central view, oblong to subcylindrical in side view, smooth under light microscope, yellowish to brownish in KOH (Fig. 2). Basidia 4-spored, (27–)27–37.4–54(–54) × (9–)9–12.3–19(–19) µm, clavate, yellowish to brownish in KOH, sterigmata up to 3 µm long. Cheilocystidia (19–)19.3–25.5–33(–35) × (4–)4.1–6–8.2(–8.5) µm, frequent, fusiform, thin-walled, yellowish to brownish hyaline in KOH and NH4OH. Pleurocystidia (40­–)41–53–69(–70) × (8–)7.4–12–16.6(–17) µm, thin-walled, fusiform to broadly fusiform with a long pedicel and sharp apex, occasionally containing yellowish inclusions, yellowish to brownish hyaline in KOH and NH4OH. Hymenophoral trama boletoid, elements smooth, cylindrical, hyaline, 5–10 µm wide. Pileipellis a trichodermium, hyphae terminations with 3–4 cells that are 5–11 µm wide and terminal cells 31–48 µm × 6–10 µm, colourless to slightly brownish in KOH. Pileus trama composed of interwoven hyaline hyphae 5–9 µm wide. Stipitipellis a disrupted hymeniderm with hyphae 3.7–7.4 µm wide, colourless to slightly brownish in KOH and caulocystidia (24–)24.5–35–47(–48) × (9–)9.2–12.4–16.9(–17) µm, thin-walled, clavate to broadly clavate with a sharp apex, yellowish to brownish hyaline in KOH and NH4OH. Stipe trama composed of cylindrical, hyaline, interwoven hyphae 3.7–7.4 µm wide. Clamp connections absent.

Figure 2.

Figure 2.

Open in a new tab

Microscopic features of Tylocinumbrevisporum; a Basidiospores; b Basidia; c, d Caulocystidia; e Pleurocystidia; f Cheilocystidia; g Pileipellis. Scale bars: a, b, c, d, f = 10 µm, e = 20 µm, g = 50 µm.

Diagnosis

This species is distinguished from Tylocinumgriseolum by its greyish-brown colour, greyish-orange to brownish-orange colour change in the hymenophore when bruised, smaller pores (≤ 0.5 mm) and longer tubes (up to 6 mm long). Additionally, the basidiospores are shorter and narrower compared to T.griseolum and the basidia are slightly longer and broader. Furthermore, the pleurocystidia of Tylocinumbrevisporum are longer than its cheilocystidia.

Etymology

Epithet “brevisporum”; from the Latin words brevi (short) and sporae (spores), referring to the shorter spores of this species compared to Tylocinumgriseolum.

Distribution

Thus far known only from northern Thailand.

Ecology

Solitary, in tropical forest dominated by Dipterocarpaceae (Dipterocarpus spp. and Shorea spp.), with some Fagaceae (Quercus spp., Lithocarpus spp. and Castanopsiscalathiformis).

Notes

Morphologically, Tylocinumbrevisporum is similar to Tylocinumgriseolum, with which it shares the overall grey colour of the basidiomata and dark scabrous stipe surface. However, Tylocinumbrevisporum is more brownish as compared to the grey Tylocinumgriseolum. In addition, Wu et al. (2016) mentioned no discolouration in the context of Tylocinumgriseolum. The context of Tylocinumbrevisporum becomes slightly brown when bruised. The hymenophore of T.brevisporum changes to greyish-orange to brownish-orange when bruised as compared to the unchanging hymenophore of T.griseolum. Moreover, T.griseolum has relatively larger pores (up to 1.5 mm) than that of T.brevisporum (< 0.5 mm). The tubes in T.griseolum are also shorter than those of T.brevisporum.

The basidiospores of Tylocinumbrevisporum [(6.7–)7.5–10–11.7(–11.8) × (3.1–)3.5–4.7–5.8(–5.9) µm, Q = (1.7–)1.79–2.15–2.5(–2.61)] are shorter and narrower than those of Tylocinumgriseolum [(11)12.0–14.5(16) × 4.5–5.5 µm Q = 2.60–3.22] from China. The basidia of T.brevisporum [(27–)27–37.4–54(–54) × (9–)9–12.3–19(–19) µm] are also slightly longer and broader than T.griseolum [30–45 × 10–12 µm]. Wu et al. (2016) reported that, for T.griseolum, the pleurocystidia and cheilocystidia are similarly-sized. In T.brevisporum, the pleurocystidia are longer than the cheilocystidia. Phylogenetically, T.brevisporum clusters with T.griseolum, together forming a well-supported clade (MLB/BPP = 93/1.00) i.e. the genus Tylocinum.

Analysis

Phylogenetic analysis

The concatenated gene dataset comprised 47 terminals. The final alignment contained 121 sequences (38 for atp6, 46 for tef1, 37 for rpb2) and was 2,676 characters long, including gaps. Both ML and Bayesian analyses produced the same tree topology; thus, only the ML tree is shown with both Maximum Likelihood Bootstrap (MLB) and Bayesian Posterior Probabilities (BPP) values. In the analyses, the new species Tylocinumbrevisporum shared a sister relationship with the type species Tylocinumgriseolum (Fig. 3), providing strong statistical support (MLB = 93 and BPP = 1.00) for the genus Tylocinum (Leccinoideae). The atp6 sequence of the holotype (BR 137) was 100% identical to OR622.

Figure 3.

Figure 3.

Open in a new tab

Maximum Likelihood phylogenetic tree inferred from the three-gene dataset (atp6, rpb2, tef1). The three Lanmaoa and three Baorangia species were used as outgroup taxa. Maximum Likelihood Bootstrap (MLB, left) ≥ 70% and Bayesian Posterior Probabilities (BPP, right) ≥ 0.95 are shown above supported branches. The new species is in bold.

Discussion

Boletales is a globally-distributed order of fungi, comprising morphologically diverse groups (Binder and Hibbett 2006, Wu et al. 2016), with ECM, ligninolytic, saprobic and mycoparasitic members (Binder and Hibbett 2006, Kirk et al. 2008). Thorough morphological and phylogenetic analyses of the order has led to the discovery of new genera and other taxa (e.g. Binder and Bresinsky 2002, Wu et al. 2014, Zhu et al. 2015, Wu et al. 2016, Orihara et al. 2016, Vadthanarat et al. 2019, Zhang et al. 2019). Boletaceae Chevall. 1826 is a morphologically diverse family currently comprising of 94 genera distributed amongst seven subfamilies (Binder and Hibbett 2006, Wu et al. 2014, Wu et al. 2016). The subfamily Leccinoideae was revealed by the phylogenetic analyses of Wu et al. (2014). Currently, this subfamily comprises fifteen genera, viz. Binderoboletus T.W. Henkel & M.E. Sm. 2016, Borofutus Hosen & Z.L. Yang 2012, Chamonixia Rolland 1899, Ionosporus Khmeln. 2018, Kaziboletus Iqbal Hosen & Zhu L. Yang 2021, Leccinum Gray 1821, Lecinellum Bresinsky & Manfr. Binder 2003, Pseudoaustroboletus Y.C. Li & Zhu L. Yang 2014, Octavania, Retiboletus Manfr. Binder & Bresinsky 2002, Rossbeevera T. Lebel & Orihara 2012, Rhodactina Pegler & T.W.K. Young 1989, Spongiforma Desjardin, Manfr. Binder, Roekring & Flegel 2009, Spongispora G. Wu, S.M.L. Lee, E. Horak & Z.L. Yang 2018, Turmalinea Orihara & N. Maek. 2015 and Tylocinum. Only ten of these genera are stipitate-pileate.

Our survey on the diversity of boletes in northern Thailand led to the discovery of a second species of Tylocinum (the focus of the present study), being found in tropical forests dominated by Dipterocarpaceae, which have been reported as ECM hosts for Boletaceae (Desjardin et al. 2009, Halling et al. 2014, Wu et al. 2018, Vadthanarat et al. 2019). According to Wu et al. (2016), the white to dirty white hymenophore of Tylocinum is similar to that of Tylopilus Karst. 1881 when young, while the verrucose stipe surface is similar to Leccinum. The stipe surface of Tylocinum is dotted-verrucose, which may give a more or less rough touch, but it does not produce markedly projecting scabers like in Leccinum. Tylocinum is also similar to Tylopilus, but there are some morphological differences between the two genera. Tylopilus species usually produce larger basidiomata and have minutely and densely tomentose to dotted-tomentose, but never dotted-verrucose, stipitipellis. Moreover, some Tylopilus species have reticulate stipe, whereas, in Tylocinum, the stipe is at most longitudinally venose near the apex. As the diversity of Boletaceae in Thailand is high and remains understudied (e.g. Vadthanarat et al. 2021), further studies may uncover additional species of Tylocinum or related taxa.

Supplementary Material

XML Treatment for Tylocinum brevisporum

Acknowledgements

B. Raghoonundon appreciates the kind support given by the Mushroom Research Foundation (MRF), Thailand. The authors thank the Thailand Science Research and Innovation (TSRI) (Grant No. DBG6280009) entitled “Macrofungi diversity research from the Lancang-Mekong Watershed and surrounding areas”. The authors are also grateful to Amy Choong for loaning her collection of Spongisporatamasekensis, Changlin Zhao for sequencing of Tylocinum specimens and to the reviewers for their comments.

References

  1. Ashton P. Dipterocarpaceae . The Hague, The Netherlands: Martinus-Nijhoff Publishers.; 1982. [Google Scholar]
  2. Binder M., Bresinsky A. Derivation of a polymorphic lineage of Gasteromycetes from boletoid ancestors. Mycologia. 2002;94:85–98. doi: 10.1080/15572536.2003.11833251. [DOI] [PubMed] [Google Scholar]
  3. Binder M, Hibbett D. S. Molecular systematics and biological diversification of Boletales. Mycologia. 2006;98:971–981. doi: 10.1080/15572536.2006.11832626. [DOI] [PubMed] [Google Scholar]
  4. Brearley F. Q. Ectomycorrhizal associations of the Dipterocarpaceae. Biotropica. 2012;44(5):637–648. doi: 10.1111/j.1744-7429.2012.00862.x. [DOI] [Google Scholar]
  5. Chuankid B, Vadthanarat S, Hyde K. D., Raspé Olivier, et al. Three new Phylloporus species from tropical China and Thailand. Mycological Progress. 2019;18(5):603–614. doi: 10.1007/s11557-019-01474-6. [DOI] [Google Scholar]
  6. Chuankid B, Vadthanarat S, Thongbai B, Stadler Marc, et al. Retiboletus (Boletaceae) in Northern Thailand: one novel species and two first records. Mycoscience. 2021;62:1–10. doi: 10.47371/mycosci.2021.05.003. [DOI] [PMC free article] [PubMed] [Google Scholar]
  7. Corrales A, Henkel T. W., Smith M. E. Ectomycorrhizal associations in the tropics–biogeography, diversity patterns and ecosystem roles. New Phytologist. 2018;220(4):1076–1091. doi: 10.1111/nph.15151. [DOI] [PubMed] [Google Scholar]
  8. Darriba D, Taboada G. L., Doallo, Posada D. jModelTest 2: more models, new heuristics and parallel computing. Nature Methods. 2012;9(8):772. doi: 10.1038/nmeth.2109. [DOI] [PMC free article] [PubMed] [Google Scholar]
  9. Desjardin D, Binder M., Roekring S., Flegel T. Spongiforma, a new genus of gasteroid boletes from Thailand. Fungal Diversity. 2009;37:1–8. [Google Scholar]
  10. Doyle J. J, Doyle JL. Isolation of plant DNA from fresh tissue. Focus. 1990;12:13–15. [Google Scholar]
  11. Halling R, Desjardin D, Fechner N, Arora D, et al. New porcini (Boletussect.Boletus) from Australia and Thailand. Mycologia. 2014;106:830–834. doi: 10.3852/13-340. [DOI] [PubMed] [Google Scholar]
  12. Hamilton R., Hall T., Stevenson J., Penny D. Distinguishing the pollen of Dipterocarpaceae from the seasonally dry and moist tropics of south-east Asia using light microscopy. Review of Palaeobotany and Palynology. 2019;263:117–133. doi: 10.1016/j.revpalbo.2019.01.012. [DOI] [Google Scholar]
  13. Hosen MI, Feng B, Wu G, Zhu XT, et al. Borofutus, a new genus of Boletaceae from tropical Asia: phylogeny, morphology and taxonomy. Fungal Diversity. 2013;58:215–226. doi: 10.1007/s13225-012-0211-8. [DOI] [Google Scholar]
  14. Jayasiri SC, Hyde KD, Ariyawansa HA, Bhat J, et al. The Faces of Fungi database: fungal names linked with morphology, phylogeny and human impacts. https://www.facesoffungi.org/ Fungal Diversity. 2015;74(1):3–18. doi: 10.1007/s13225-015-0351-8. [DOI] [Google Scholar]
  15. Katoh K, Standley DM. MAFFT Multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution. 2013;30(4):772–780. doi: 10.1093/molbev/mst010. [DOI] [PMC free article] [PubMed] [Google Scholar]
  16. Khmelnitsky O, Davoodian N, Singh P, et al. Ionosporus: a new genus for Boletuslongipes (Boletaceae), with a new species, I.australis, from Australia. Mycological Progress. 2019;18(3):439–451. doi: 10.1007/s11557-018-01463-1. [DOI] [Google Scholar]
  17. Kirk P. M., Cannon P. F., Minter D. W., Stalpers J. A. Dictionary of the Fungi. 10th edition. CABI International, Wallingford.; 2008. [Google Scholar]
  18. Kornerup A, Wanscher J. H. Methuen Handbook of Colour. Eyre Methuen; 1978. [Google Scholar]
  19. Matheny P. B. Improving phylogenetic inference of mushrooms with RPB1 and RPB2 nucleotide sequences (Inocybe; Agaricales). Molecular Phylogenetics and Evolution. 2005;35:1–20. doi: 10.1016/j.ympev.2004.11.014. [DOI] [PubMed] [Google Scholar]
  20. Miller Mark A., Pfeiffer W, Schwartz T. The CIPRES science gateway: enabling high-impact science for phylogenetics researchers with limited resources. XSEDE '12: Proceedings of the 1st Conference of the Extreme Science and Engineering Discovery Environment: Bridging from the eXtreme to the campus and beyond. 2012;39:1–8. doi: 10.1145/2335755.2335836. [DOI] [Google Scholar]
  21. Neves M. A., Binder M., Halling R., Hibbett D, et al. The phylogeny of selected Phylloporus species inferred from NUC–LSU and ITS sequences, and descriptions of new species from the Old World. Fungal Diversity. 2012;55(1):109–123. doi: 10.1007/s13225-012-0154-0. [DOI] [Google Scholar]
  22. Nuhn M. E., Binder M., Taylor A. F.S, Halling R. E, et al. Phylogenetic overview of the Boletineae. Fungal Biology. 2013;117:479–511. doi: 10.1016/j.funbio.2013.04.008. [DOI] [PubMed] [Google Scholar]
  23. Orihara T, Smith M. E, Shimomura N., Iwase K, et al. Diversity and systematics of the sequestrate genus Octaviania in Japan: two new subgenera and eleven new species. Persoonia. 2012;28:85–112. doi: 10.3767/003158512X650121. [DOI] [PMC free article] [PubMed] [Google Scholar]
  24. Orihara T., Lebel T., Ge Z. W., Smith M. E, et al. Evolutionary history of the sequestrate genus Rossbeevera (Boletaceae) reveals a new genus Turmalinea and highlights the utility of ITS minisatellite–like insertions for molecular identification. Persoonia. 2016;37:173–198. doi: 10.3767/003158516X691212. [DOI] [PMC free article] [PubMed] [Google Scholar]
  25. Phookamsak R, Hyde K. D, Jeewon R, et al. Fungal diversity notes 929-1035: taxonomic and phylogenetic contributions on genera and species of fungi. Fungal Diversity. 2019;95:1–273. doi: 10.1007/s13225-019-00421-w. [DOI] [Google Scholar]
  26. Raspé O., Vadthanarat S., Kesel A. De, Degreef J., et al. Pulveroboletusfragrans, a new Boletaceae species from Northern Thailand, with a remarkable aromatic odor. Mycological Progress. 2016;15(38):38. doi: 10.1007/s11557-016-1179-7. [DOI] [Google Scholar]
  27. Rehner S. A, Buckley E. A Beauveria phylogeny inferred from nuclear ITS and EF1-alpha sequences: evidence for cryptic diversification and links to Cordyceps teleomorphs. Mycologia. 2005;97:84–98. doi: 10.1080/15572536.2006.11832842. [DOI] [PubMed] [Google Scholar]
  28. Ronquist F, Teslenko M, Mark P Van Der, Ayres DL, et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology. 2012;61(3):539–542. doi: 10.1093/sysbio/sys029. [DOI] [PMC free article] [PubMed] [Google Scholar]
  29. Smith S. E, Read D. J, Smith S. E, Read D. Structure and development of ectomycorrhizal roots. Mycorrhizal Symbiosis. 2008;3:191–268. doi: 10.1002/9781118951446.ch4. [DOI] [Google Scholar]
  30. Stamatakis A. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics. 2006;22(21):2688–2690. doi: 10.1093/bioinformatics/btl446. [DOI] [PubMed] [Google Scholar]
  31. Vadthanarat S, Raspé O, Lumyong S. Phylogenetic affinities of the sequestrate genus Rhodactina (Boletaceae), with a new species, R.rostratispora from Thailand. MycoKeys. 2018;29:63–80. doi: 10.3897/mycokeys.29.22572. [DOI] [PMC free article] [PubMed] [Google Scholar]
  32. Vadthanarat S, Saisamorn L., Raspé O. Cacaoporus, a new Boletaceae genus, with two new species from Thailand. MycoKeys. 2019;54:1–29. doi: 10.3897/mycokeys.54.35018. [DOI] [PMC free article] [PubMed] [Google Scholar]
  33. Vadthanarat S, Halling R. E, Amalfi M., Lumyong S. An unexpectedly high number of new Sutorius (Boletaceae) species from Northern and Northeastern Thailand. Frontiers in Microbiology. 2021;12:643505. doi: 10.3389/fmicb.2021.643505. [DOI] [PMC free article] [PubMed] [Google Scholar]
  34. Watling R, Lee S. S, Turnbull E. The occurrence and distribution of putative ectomycorrhizal basidiomycetes in a regenerating South-East Asian rain forest. Tropical Mycology. 2002;1:25–43. [Google Scholar]
  35. Wu G, Feng B, Xu J, et al. Molecular phylogenetic analyses redefine seven major clades and reveal 22 new generic clades in the fungal family Boletaceae. Fungal Diversity. 2014;69:93–115. doi: 10.1007/s13225-014-0283-8. [DOI] [Google Scholar]
  36. Wu G, Zhao K, Li Y. C, et al. Four new genera of the fungal family Boletaceae. Fungal Diversity. 2015;81:1–24. doi: 10.1007/s13225-015-0322-0. [DOI] [Google Scholar]
  37. Wu G, Li Y. C., Zhu X. T. One hundred noteworthy boletes from China. Fungal Diversity. 2016;81:25–188. doi: 10.1007/s13225-016-0375-8. [DOI] [Google Scholar]
  38. Wu G, Lee S. M.L, Horak E, Yang Z. L. Spongisporatemakensis, a new boletoid genus and species from Singapore. Mycologia. 2018;1105:919–929. doi: 10.1080/00275514.2018.1496387. [DOI] [PubMed] [Google Scholar]
  39. Yuwa-Amornpitak T., Vichitsoonthonkul T., Tanticharoen M., Cheevadhanarak S., et al. Diversity of ectomycorrhizal fungi on Dipterocarpaceae in Thailand. Journal of Biological Sciences. 2006;6:1059–1064. doi: 10.3923/jbs.2006.1059.1064. [DOI] [Google Scholar]
  40. Zhang M., Li T. H., Wang C. Q., Zeng N. K., et al. Phylogenetic overview of Aureoboletus (Boletaceae, Boletales), with descriptions of six new species from China. MycoKeys. 2019;61:111–145. doi: 10.3897/mycokeys.61.47520. [DOI] [PMC free article] [PubMed] [Google Scholar]
  41. Zhu X. T, Wu G, Zhao K., Halling R. E, et al. Hourangia, a new genus of Boletaceae to accommodate Xerocomuscheoi and its allied species. Mycological Progress. 2015;14:37. doi: 10.1007/s11557-015-1060-0. [DOI] [Google Scholar]

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Tylocinum brevisporum
BDJ.9.e75907-treatment1.xml (25.5KB, xml)

Articles from Biodiversity Data Journal are provided here courtesy of Pensoft Publishers

RESOURCES