Research Article |
Corresponding author: Olivier Raspé ( Olivier.Ras@mfu.ac.th ) Academic editor: Rui-Lin Zhao
© 2023 Monthien Phonemany, Santhiti Vadthanarat, Bhavesh Raghoonundon, Naritsada Thongklang, 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:
Phonemany M, Vadthanarat S, Raghoonundon B, Thongklang N, Raspé O (2023) Additions to Hohenbuehelia (Basidiomycota, Pleurotaceae): two new species and notes on H. tristis from northern Thailand. MycoKeys 99: 109-130. https://doi.org/10.3897/mycokeys.99.105317
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Two new species and a first geographical record of Hohenbuehelia are described from Thailand. Macroscopic and microscopic descriptions with photoplates, as well as a multigene phylogeny are provided. Hohenbuehelia flabelliformis sp. nov. is recognised by large flabelliform basidiomata, densely villose yellowish-white pileus with white hairs near the point of attachment, basidiospores that mostly are ellipsoid in front view and phaseoliform in side view, the absence of cheilocystidia, and a trichoderm pileipellis. Hohenbuehelia lageniformis sp. nov. is characterised by fleshy basidiomata, velutinous pileus with whitish hairs near the point of attachment and the margin, elsewhere pale greyish-yellow and with only sparse white hairs, pale brown to light brown and mucilaginous context, subglobose basidiospores, lageniform cheilocystidia, an ixotrichoderm pileipellis, and the absence of pileoleptocystidia. Hohenbuehelia tristis is characterised by small creamy-white, spathuliform basidiomata that are larger than the type subspecies, minutely pubescent pileus with tiny greyish hairs that disappear when mature, leaving the surface glutinous, faintly translucent and shiny, ellipsoid to sub-ellipsoid basidiospores, lecythiform to sublageniform cheilocystidia, and an ixotrichoderm pileipellis. Hohenbuehelia tristis is recorded for Thailand for the first time. Based on the polymorphism observed in part of the nrLSU gene, the presence of two divergent lineages within H. tristis is discussed.
Agaricales, DNA sequence heteromorphisms, molecular phylogeny, pleurotoid mushrooms, Southeast Asia, taxonomy, two new species
Hohenbuehelia Schulzer belongs to the family Pleurotaceae Kühner in the order Agaricales Underw. In former studies, the asexual stages of Hohenbuehelia species were separately placed in the genus Nematoctonus Drechsler (
A few Hohenbuehelia species have been reported as edible. However, they have very low culinary value (
Thailand has a high mushroom diversity with many new species yet to be discovered (
The mushroom specimens were collected during the rainy season in 2019 and 2020 from Chiang Mai and Chiang Rai Provinces, in northern Thailand. The fresh basidiomata were photographed in situ. Details including collecting date, locality, habitat and ecology of the surroundings, were noted. The specimens were wrapped in aluminium foil or kept in plastic boxes and brought back to the lab for morphological descriptions.
Macromorphological descriptions were done, based on the fresh specimens and colour codes were given following the colour charts of
Genomic DNA was extracted from the dried herbarium specimens using the Biospin Fungus Genomic DNA Extraction Kit (Bioer Technology, Hangzhou, China), following the manufacturer’s instructions. The ITS region and parts of the nrLSU and tef1 genes were amplified by a polymerase chain reaction (PCR) and sequenced. The following primers were used: ITS1-F and ITS4 for ITS (
Sequence reads were checked using Bioedit Sequence Alignment Editor version 7.0.9.0 and assembled using SeqMan (DNAstar, Madison, WI, USA). Each sequence was blasted using the Basic Local Alignment Search Tool (BLAST) against the National Center for Biotechnology Information (NCBI) database (http://www.ncbi.nlm.nih.gov/genbank/) to check that it was from the correct genus and not from contamination, as well as to find the closest matches. Newly-generated sequences were deposited in GenBank. All sequences (Table
GenBank accession numbers and geographical origins of taxa used in the phylogenetic analyses.
Species | Specimen/culture | Country | GenBank accession numbers | ||
---|---|---|---|---|---|
ITS | nLSU | tef1 | |||
Hohenbuehelia algonquinensis | RGT 870601/12 UWO (culture T-434) | Canada | KU355341 | AF139950 | KU355456 |
H. angustata | CBS 856.85 | Canada | MH861919 | MH873608 | – |
H. atrocoerulea | AMB 18080 | Hungary | KU355304 | KU355389 | KU355439 |
H. auriscalpium | PRMJH1372007 | Czech Republic | MT525862 | MT534054 | – |
H. bonii | K(M):165700 | England | KX064444 | – | – |
H. boullardii | JCM06005 | Spain | MG553637 | MG553644 | – |
H. canadensis, as H. atrocoerulea var. grisea | DAOM 158848 | Canada | KU355356 | – | – |
H. carlothornii | AMB 18106 | Costa Rica | KY698012 | KY698013 | – |
H. chevallieri | WU:6528 | Austria | KT388040 | – | – |
H. culmicola | Roux 3488 | Italy | KU355323 | – | – |
H. cyphelliformis | Z+ZT 994 | Switzerland | KU355325 | KU355393 | KU355445 |
H. faerberioides | Mertens | France | MG553638 | MG553645 | MW240984 |
H. flabelliformis | MFLU22-0008 | Thailand | OP236779 | OM521957 | OM714821 |
H. flabelliformis | MFLU22-0009 | Thailand | OP236780 | – | – |
H. fluxilis | WU 29608 | Austria | KU355326 | – | – |
H. grisea | MCVE 27293, as H. myxotricha | Italy | KU355329 | KU355394 | KU355447 |
H. grisea | VPI-F-0001921 (culture VT 1324 = T-132), as H. nigra | USA | KY679143 | KY679143 | – |
H. grisea | MFLUCC 12-0451 | Thailand | MF150036 | – | – |
H. grisea | HFJAU0029 | China | MN258645 | – | – |
H. ilerdensis | Roux 3924 | Spain | MG553639 | MG553646 | |
H. josserandii | P.K. EG10-812-T-F | Germany | KU355353 | KU355403 | KU355463 |
H. lageniformis | MFLU22-0010 | Thailand | OP236781 | OM521958 | OM763737 |
H. lageniformis | MFLU22-0012 | Thailand | OP236783 | OM521959 | – |
H. leightonii | WU 5846 | Spain | MG553640 | MG553647 | – |
H. ligulata | PDD:80775 | New Zealand | KM975439 | – | – |
H. longipes | LIP 0400317 | Italy | KU355333 | KU355396 | KU355449 |
H. mastrucata | TRTC 152314 | Italy | KU355336 | KU355397 | KU355451 |
H. mustialensis | DAOM 46374 | Canada | KY124252 | – | – |
H. nimueae | RGT 871128/01 UWO (culture T-489 = CBS 212.91), as H. nigra | Canada | KY679144 | KY679144 | – |
H. odorata | TBGT17443 | India | MN059651 | – | – |
H. petaloides | AMB 18088 | Italy | KU355346 | KU355402 | KU355460 |
H. pinacearum | DAOM 84313 | Canada | MH137814 | MH137837 | – |
H. portegna | J.E. Wright 1136 BAFC | Argentina | AF139959 | AF139959 | – |
H. tristis | MFLU22-0015 | Thailand | OP355451 | OM521961 | – |
H. tristis | MFLU22-0016 | Thailand | OP355450 | OM521962 | ON394004 |
H. reniformis | HMJAU7091 | China | GQ142024 | GQ142041 | – |
H. robusta | CBS 130.68 | – | MH859087 | MH870800 | – |
H. thornii | WU 21790 | Portugal | KU355343 | KU355401 | KU355457 |
H. tremula | M 0223665 | Italy | KU355358 | KU355406 | KU355466 |
H. tristis | RV95/214 DUKE | Australia | – | AF042601 | – |
H. tristis | RV95/295 DUKE | Australia | – | AF135171 | – |
H. unguicularis | Z+ZT 1112 | France | KU355361 | KU355408 | KU355467 |
H. valesiaca | Roux 2975 | France | KU355340 | KU355399 | KU355455 |
H. wilhelmii | Z+ZT n. 1154 | France | MF494947 | MF494948 | MF494949 |
Pleurotus citrinopileatus | YAASM1585 | China | KX836372 | – | KX840311 |
P. eryngii | CCMSSC00480 | China | KX836350 | – | KX840226 |
P. ostreatus | TENN 53662 (= AFTOL-ID 564) | Austria | AY854077 | AY645052 | AY883432 |
P. djamor, as P. placentodes | HKAS51745 | China | KR827693 | KR827695 | KR827699 |
P. pulmonarius | BCRC36906 | Taiwan | MH453616 | MH447275 | MH500353 |
All phylogenetic analyses were done on the CIPRES Science Gateway version 3.3 web server (
The combined dataset consisted of 39 Hohenbuehelia and five Pleurotus accessions (Table
Phylogeny of selected sequences of Hohenbuehelia based on a Maximum Likelihood analysis of three nuclear gene regions (nrITS, nrLSU, tef1). The Maximum Likelihood bootstrap values (BS ≥ 70%) and Bayesian posterior probabilities values (PP ≥ 0.90) are shown on the branches. Newly-sequenced collections are in bold. Five Pleurotus species were used as outgroup. (T) designates holotypes. The sequence H. flabelliformis voucher number MFLU22-0008 was identical to MFLU22-0009 (ITS only). Sequences of H. lageniformis voucher number MFLU22-0010 were identical to MFLU22-0012 (ITS and LSU). Sequences of H. tristis voucher number MFLU22-0016 were identical to MFLU22-0015 (ITS and LSU) and to the two specimens MFLUCC 12-0451 and HFJAU0029, identified as H. grisea (both only ITS) from GenBank, except for three substitution heteromorphisms in the ITS sequence of MFLU22-0016 (see Table
Positions of substitution heteromorphisms in the ITS sequence of H. tristis MFLU22-0016 and corresponding character states in other H. tristis ITS sequences.
This species is distinguished from other Hohenbuehelia species by large flabelliform basidiomata, yellowish-white pileus that is densely villose with white hairs longer near the point of attachment, and shorter towards the margin, ellipsoid basidiospores, absence of cheilocystidia, and a trichoderm pileipellis.
Thailand, Chiang Mai Province, Mae Taeng District, Pha Daeng Village, 27 May 2019, Monthien Phonemany (MFLU22-0008).
"flabelliformis" refers to the flabelliform shape of basidiomata.
Pileus 35–45 × 20–40 mm, spathulate when young, expanding to spathuliform, flabelliform, rounded flabelliform or orbicular, white when young, becoming yellowish-white to pale yellow (4A2–4A3) at the centre and cream (2A3–2B3) to slightly darker elsewhere in age; surface densely villose with white hairs that are longer near the attachment and shorter towards the margin, as observed with a lens; margin white, incurved when young, becoming straight when old. Lamellae 1–3 mm wide, decurrent, pale yellow to yellowish-white (4A3–4A2), moderately crowded when mature, with lamellulae in 1–3 tiers; edge concolorous to sides, fimbriate. Stipe absent or as pseudostipe 5–12 mm × 3–8 mm. Context consisting of two layers: 1) non-gelatinous layer, 1 mm thick, soft when young and rather leathery when old, white to dirty white (4A1–4A2); 2) gelatinous layer 0.5 mm thick, soft, sticky, colourless. Odour mild, pleasant. Taste none. Spore print white.
Basidiospores [150/3/2] (5.8–)6–7–8(–8.6) × (3.5–)4–4.2–5(–5.1) µm, Q = (1.3–)1.36–1.67–2.01(–2.03), ellipsoid to elongate (oblong) in side view, smooth, thin-walled, inamyloid. Basidia (21–)21–25.8–35(–37) × (5–)5.3–7.3–10.5(–11) µm, subclavate to clavate, with 4 sterigmata 4–8 µm long, hyaline, smooth, thin-walled. Cheilocystidia absent. Pleurocystidia metuloidal, present on both sides of lamellae and visible with a lens, (34–)34–42–54(–55) × (8.5–)8.5–11–14(–14.5) µm, scattered, narrowly fusiform to fusiform, mucronate at apex, brownish in KOH. Hymenophoral trama irregular, hyphae 2–5 µm wide. Pileipellis a trichoderm, hyaline in KOH, brownish in water, with cylindrical terminal elements 34–86 × 4–7 µm. Pileoleptocystidia absent. Pileus trama consists of two different layers: 1) upper layer gelatinous, composed of horizontally arranged, smooth, colourless encrusted hyphae, 2–5 µm wide; 2) lower layer, non-gelatinous, composed of interwoven, smooth, hyaline hyphae, 2–4 µm wide. Clamp connections present in pileipellis, pileus trama, and hymenophoral trama.
On dead wood, scattered or fasciculate by 2–4 basidiomata. So far only found in tropical forests of northern Thailand.
Thailand. Chiang Rai Province, Pa Daed District, Pa Ngae Village, 9 August 2019, Monthien Phonemany (MFLU22-0009).
The basidiomata colour of Hohenbuehelia flabelliformis is similar to H. angustata (Berk.) Singer, H. bonii A.M. Ainsw., H. concentrica Corner, H. carlothornii Consiglio, Setti & Thorn, H. horrida (Boedijn) Corner, H. luteola G. Stev, H. malesiana Corner, H. odorata C.K. Pradeep & Bijeesh, H. olivacea Yu Liu & T. Bau, and H. testudo (Berk.) Pegler. The basidiomata range from white, yellowish-white, yellow-brown, to pinkish-orange, and are spathulate to flabelliform. Hohenbuehelia angustata, originally described from Brazil, differs from H. flabelliformis by its smaller, smooth, greyish-yellow basidiomata, with stipe 4.5 mm long, smaller basidiospores (3.5–5 × 2.5–3.5 µm), smaller cheilocystidia, and the presence of pileocystidia (
Phylogenetically, H. flabelliformis was closely related to H. algonquinensis Consiglio, Setti & Thorn. (voucher RGT 870601/12 UWO) with 3.5% (21/599) differences in the ITS sequence, 1.4% (12/839) in the LSU sequence, and 6.66% (35/540) in the tef1 sequence. Moreover, the morphologies of both species are completely different, with H. algonquinensis having glossy black pileus, ungulate to dimidiate basidiomata and contrasting white or off-white lamellae (
This species is distinguished from other Hohenbuehelia species by having velutinous pileus with whitish hairs near the point of attachment, and at the margin, elsewhere pale greyish-yellow, and with only sparse white hairs, pale brown to light brown and mucilaginous context, subglobose to ellipsoid basidiospores, lageniform cheilocystidia, an ixotrichoderm pileipellis, and the absence of pileoleptocystidia.
Thailand, Chiang Mai Province, Mae On District, Huai Kaew Subdistrict, Pok Village, 29 June 2019, Monthien Phonemany (MFLU22-0010).
"lageniformis" refers to the lageniform shape of cheilocystidia.
Pileus 25–30 × 15–25 mm, spathuliform when young, rounded flabelliform to sub-rounded flabelliform when mature, light yellow (4A4), pale orange to orange white (5A2–5A3), greyish-orange (5B4) becoming darker with age, sometimes sessile or with laterally attached stipe; surface moist, shiny, velutinous with whitish (1A1) hairs near the point of attachment, elsewhere light yellow to pale greyish-yellow (4A4–4B4); margin discolorous from pileus, velutinous with whitish hairs as observed under the lens, incurved even when mature, entire or sometimes undulate when old. Lamellae 1–2 mm wide, decurrent, white to pale orange (5A1–5A2) becoming slightly dark in age, close to moderately crowded, in 2–3 tiers. Stipe absent or sometimes with pseudostipe 0.5–1 mm long when young and disappearing when mature. Context consisting of two layers: 1) non-gelatinous layer, 1 mm thick, fleshy, white (1A1); 2) gelatinous layer, 1–2 mm thick, soft, sticky, brownish-orange to light brown (5C5–5D5). Odour mild. Taste none. Spore print white.
Basidiospores [150/3/3] (6.9–)7–8.8–10(–10.5) × (5.5–)6–7.0–8(–8.3) µm, Q = (0.99–)1.11–1.26–1.49(–1.53) subglobose to ellipsoid in front view, ellipsoid to phaseoliform in side view, smooth, thin-walled, inamyloid. Basidia (24.3–)24–31.9–53(–58.2) × (5.4–)5.5–9.4–14(–14.6) µm, clavate, with (2)–4 sterigmata, 4–8 µm long, hyaline, smooth, thin-walled. Cheilocystidia (17.6–)18–21.9–26.5(–26.9) × (3.2–)3–5.0–9(–8.9) µm, lageniform with an inflated base with a thin, rostrate apex, hyaline, thin-walled. Pleurocystidia (31–)31–50.3–70(–71) × (9.7–)10–15.3–20(–20.3) µm, metuloidal, setiform, narrowly fusiform to fusiform, encrusted with crystals, brownish or yellowish in water, colourless, but still encrusted with crystals in KOH. Hymenophoral trama subregular, hyphae 2–4 µm wide. Pileipellis an intricate trichoderm with cylindrical terminal elements 41–122 × 2–5 µm. Pileoleptocystidia absent. Pileus trama consisting of two layers: 1) upper layer gelatinous, composed horizontally arranged, smooth, colourless, encrusted hyphae, 1–4 µm wide; 2) non-gelatinous layer, composed interwoven, smooth with hyaline hyphae, 2–5 µm wide. Clamp connections present in pileipellis, pileus trama and hymenophoral trama.
Solitary, gregarious to imbricate, on decaying branches in a tropical forests in northern Thailand.
Thailand. Chiang Rai Province, Muang Chiang Rai District, Mae Yao Subdistrict, Huai Mae Sai Village, 10 July 2019, Monthien Phonemany (MFLU22-0011); ibid., 10 July 2019, Monthien Phonemany (MFLU22-0012); ibid., 10 July 2019, Monthien Phonemany (MFLU22-0013); Pa Daed District, Pa Ngae Village, 7 September 2019, Monthien Phonemany (MFLU22-0014).
Hohenbuehelia lageniformis is characterised by velutinous pileus with whitish hairs near the point of attachment and at the margin, elsewhere pale greyish-yellow and with only sparse white hairs, subglobose to ellipsoid basidiospores, and the absence of pileoleptocystidia. Hohenbuehelia angustata, H. bonii, H. carlothornii, H. concentrica, H. flabelliformis, H. horrida, H. luteola, H. mellea Corner, and H. odorata have similar pileus colour, for example, pale orange to orange white, light yellow, yellowish-white, yellow-brown, honey-yellow or ochraceous brownish. However, those species have significant morphological differences. Hohenbuehelia angustata differs from H. lageniformis by having a smooth pileus surface, serrate margin, smaller basidiospores (3.5–5 × 2.5–3.5 µm), cutis pileipellis (
In the phylogenic tree, the most closely-related species to H. lageniformis was H. odorata (voucher TBGT17443). However, the genetic distance between the ITS sequence of H. lageniformis and H. odorata was 4.62% (27/584), which supports the distinction of the two species. Moreover, these two species also show morphological differences (see above).
The following description is based solely on the Thai materials we collected and examined.
Pileus 15–20 × 20–30 mm, spathuliform to reniform, dimidiate to orbicular, greyish-white (1B1) to yellowish-white (1B2) when young, becoming white (1A1) in age, glutinous, very faintly translucent, shiny when mature; upper surface minutely pubescent with greyish hairs (5A1–5B1) near the point of attachment and more sparsely so towards the margin as observed with a lens, with hairs disappearing in age; margin incurved becoming upturned in age. Lamellae radiating from point of attachment, 1 mm broad, very crowded, white (1A1) to pale yellow (1A3); lamellulae in 1–4 tiers. Stipe absent or pseudostipe sometimes present, laterally or dorsally attached, 1 mm long when young, then disappearing when old. Context consisting of two layers: 1) leathery layer, 1 mm thick; 2) gelatinous layer, 1–2 mm thick, soft, sticky, colourless. Odour and taste not observed. Spore print white.
Basidiospores [150/3/2] (5.1–)6–6.8–8(–9) × (3.5–)3.5–4.0–5(–5.2), Q = (1.21–)1.38–1.70–2.08(–2.26), ellipsoid, sub-ellipsoid to elongate, smooth, thin-walled, inamyloid. Basidia (13–)14–21.2–24(–24.2) × (5–)5.5–6.6–7(–7) µm, clavate to subcylindrical, mostly with four sometimes with two sterigmata, 2–4 µm long, hyaline, smooth, thin-walled. Cheilocystidia (10.9–)11–13.1–19(–19.8) × (4.2–)4.4–5.5–6.5(–6.5) µm, lecythiform to sublageniform, with subcapitate to capitate apex, hyaline, thin-walled. Pleurocystidia (38–)38–61.5–82(–82) × (10.7–)11–15.0–18.5(–18.6) µm, subfusiform, narrowly fusiform to fusiform, brownish, encrusted with crystals when observed in water, but crystals disappearing in KOH. Hymenophoral trama subregular, hyphae 2–4 µm wide. Pileipellis as tufts of ixotrichoderm with cylindrical terminal elements 40–105 × 3–5.5 µm, with light brown intracellular pigments. Pileoleptocystidia absent. Pileus trama consisting of two layers: 1) upper layer gelatinous, composed of horizontally arranged, smooth, colourless, encrusted hyphae, 1.5–2.5 µm wide; 2) lower layer non-gelatinous, composed of interwoven, smooth and hyaline hyphae, width 2–4 µm. Clamp connections present in pileipellis, pileus trama and hymenophoral trama.
Solitary, gregarious to imbricate, on dead small branches.
Thailand. Chiang Rai Province, Muang Chiang Rai District, Nang Lae Nai Village, 31 July 2019, Monthien Phonemany (MFLU22-0015). Chiang Rai Province, Muang Chiang Rai District, Mae Yao subdistrict, Huai Mae Sai Village, 31 July 2019, Monthien Phonemany (MFLU22-0016).
Two accessions identified as H. grisea, the culture MFLUCC 12-0451 from Thailand and HFJAU0029 from China (unpublished), had the same ITS sequence than H. tristis (MFLU22-0015 and MFLU22-0016) except for three substitution heteromorphisms in the ITS sequence of MFLU22-0016 (see Table
Position of substitutions and substitution heteromorphisms in the LSU sequence of H. tristis MFLU22-0015, MFLU22-0016, and corresponding character states in H. tristis RV95/214 DUKE, RV95/295 DUKE.
Genetic distance (number of substitutions, excluding heteromorphisms) between LSU sequences of closely-related Hohenbuehelia species.
Species name/ specimen number | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 |
---|---|---|---|---|---|---|---|---|---|
1. H. tristis MFLU22-0015 | |||||||||
2. H. tristis MFLU22-0016 | 0 | ||||||||
3. H. atrocoerulea AMB 18080 | 13 | 13 | |||||||
4. H. carlothornii AMB 18106 | 3 | 3 | 14 | ||||||
5. H. reniformis HMJAU7091 | 11 | 11 | 2 | 12 | |||||
6. H. robusta CBS 130.68 | 3 | 3 | 14 | 0 | 12 | ||||
7. H. tristis RV95/214 DUKE | 3 | 2 | 15 | 5 | 13 | 5 | |||
8. H. tristis RV95/295 DUKE | 3 | 1 | 14 | 4 | 12 | 4 | 2 | ||
9. H. longipes LIP 0400317 | 24 | 24 | 15 | 23 | 13 | 23 | 24 | 23 | |
10. H. tremula M 0223665 | 25 | 25 | 16 | 24 | 14 | 24 | 25 | 24 | 1 |
The Pleurotaceae belong to the Agaricales and comprise the monophyletic pleurotoid genera Pleurotus and Hohenbuehelia (
Some of the recently-described species were still introduced, based on only single-gene molecular evidence. In this study, we provide multiple-gene sequence data and detailed descriptions supporting the introduction of two new Hohenbuehelia species and a note on H. tristis from Thailand. At present, a total of six Hohenbuehelia species have been reported from Thailand including the ones in this study and three that were previously reported, namely H. panelloides, H. petaloides, and H. reniformis (
Monthien phonemany appreciates the kind support from a Thesis Writing Grant of Mae Fah Luang University, Thailand. The support from the Thailand Research Fund (TRF) grant "Study of saprobic Agaricales in Thailand to find new industrial mushroom products" (Grant No. DBG6180015) and the Thailand Science Research and Innovation (TSRI) grant "Macrofungi diversity research from the Lancang-Mekong Watershed and surrounding areas" (grant No. DBG6280009) is also acknowledged.
The authors have declared that no competing interests exist.
No ethical statement was reported.
Thailand Research Fund and Thailand Science Research and Innovation.
Conceptualization: OR, MP. Formal analysis: MP. Funding acquisition: NT. Investigation: MP. Methodology: OR. Project administration: NT. Supervision: OR. Validation: SV. Writing – original draft: MP, OR. Writing – review and editing: BR, SV, NT, OR.
Monthien Phonemany https://orcid.org/0000-0002-7865-4659
Santhiti Vadthanarat https://orcid.org/0000-0002-9035-0375
Bhavesh Raghoonundon https://orcid.org/0000-0001-6671-2404
Naritsada Thongklang https://orcid.org/0000-0001-9337-5001
Olivier Raspé https://orcid.org/0000-0002-8426-2133
All of the data that support the findings of this study are available in the main text, or in publicly accessible data repositories, as indicated in the text.