69
Th ee new species and one new eco d o Scle ode ma
(Scle ode ma aceae, Bole ales) om no he n Thailand
Didsanu da Gonkhom1,2 , Phongeun Sysouphan hong2, Ma c S adle 3,4 , Na i sada Thongklang1,2 ,
Ke in D. Hyde2
1 School o Science, Mae Fah Luang Uni e si y, Chiang Rai, Thailand
2 Cen e o Excellence in Fungal Resea ch, Mae Fah Luang Uni e si y, Chiang Rai, Thailand
3 Depa men o Mic obial D ugs, Helmhol z Cen e o In ec ion Resea ch (HZI), 38124 B aunschweig, Ge many
4 Ge man Cen e o In ec ion Resea ch (DZIF), pa ne si e Hanno e /B aunschweig, Inho ens asse 7, 38124 B aunschweig, Ge many
Co esponding au ho s: Na i sada Thongklang ([email p o ec ed]); Ke in D. Hyde ([email p o ec ed])
Copy igh : © Didsanu da Gonkhom e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
The genus Scle ode ma (Scle ode ma aceae) con ains gas e oid ec omyco hizal ungi
and is dis ibu ed wo ldwide in empe a e and opical egions. F esh specimens we e
collec ed in Thailand and epo h ee undesc ibed species and one new eco d o he
coun y. These species we e cha ac e ized by pho og aphs o eshly collec ed basidi-
omes, and hei mac o- and mic oscopic ea u es we e compa ed wi h hose o known
species o Scle ode ma. Addi ionally, DNA sequence da a we e gene a ed o ou loci,
including he nuclea ibosomal in e nal ansc ibed space egion (ITS), he la ge sub-
uni ibosomal RNA gene (LSU), he ansla ion elonga ion ac o 1-alpha gene ( e 1-α),
and he second la ges subuni o RNA polyme ase II ( pb2). A mul i-locus phylogeny
was cons uc ed o con i m hei axonomic placemen .
Key wo ds: Basidiomes, Bole ales, mo phology, axonomy, h ee new species
In oduc ion
Ec omyco hizal ungi o m symbio ic ela ionships wi h he eede oo s o
many ee species and o he plan s ha bene i bo h pa ne s (Cha ya and
Ga g 2019). They a e dis ibu ed wo ldwide in empe a e and opical egions
(Co ales e al. 2018). The ec omyco hizal ungi include he genus Scle o-
de ma (Oua iki e al. 2022). Pe soon (1801) e ec ed he genus Scle ode ma,
which was la e upda ed by Guzmán (1970), who p oposed in agene ic
classi ica ions such as sec ions Scle angium, Scle ode ma, and Mac ospo a
based on mo phology. Mode n phylogene ic s udies ha e u he e ined
hese g oupings (de Menezes Filho e al. 2022; Wu e al. 2023). F om ea ly
mo phology-based classi ica ions o mo e ecen gene ic in es iga ions ha
ha e imp o ed ou unde s anding o species ela ionships (Pe soon 1801;
Guzmán 1970; Wu e al. 2023), he his o y o Scle ode ma demons a es he
e ol ing dynamics o ungal axonomy. The e olu ion om Lycope don e u-
cosum o S. e ucosum illus a es how scien i ic classi ica ion con inues o
e ol e (Pe soon 1801).
Academic edi o : Tho s en Lumbsch
Recei ed:
28 May 2025
Accep ed:
26 Augus 2025
Published:
8 Oc obe 2025
Ci a ion: Gonkhom D, Sysouphan hong
P, S adle M, Thongklang N, Hyde
KD (2025) Th ee new species and
one new eco d o Scle ode ma
(Scle ode ma aceae, Bole ales) om
no he n Thailand. MycoKeys 123:
69–88. h ps://doi.o g/10.3897/
mycokeys.123.160438
MycoKeys 123: 69–88 (2025)
DOI: 10.3897/mycokeys.123.160438
70
MycoKeys 123: 69–88 (2025), DOI: 10.3897/mycokeys.123.160438
Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
The e a e 206 Scle ode ma species lis ed in Index Fungo um (h ps://www.
index ungo um.o g/Names/Names.asp, accessed on 24 Ma ch 2025) and 76
species in Species Fungo um (h ps://www.species ungo um.o g/Names/
Names.asp, accessed on 24 Ma ch 2025). The genus belongs o he amily
Scle ode ma aceae, o de Bole ales, and class Aga icomyce es (Binde and
Hibbe 2006).
Scle ode ma species ha e adi ionally been seg ega ed acco ding o he
mo phology o he basidioma a and he su ace o he pe idium, he ype o
dehiscence o he pe idium, he colo o he gleba, and he o namen a ion o
hei basidiospo es (de Menezes Filho e al. 2022). Mo eo e , he hickness
and scaliness o he pe idium, he p esence o s alks o he basidiome, and
he o m o he s ipe ha e occasionally been used o dis inguish species in he
genus (Rau e al. 2020).
Al hough egions such as Eu ope and Ame ica a e ela i ely well s udied in
e ms o Scle ode ma axonomy, da a a e lacking o opical A ica (Sanon
e al. 1997) and Asia (Fa me and Syl ia 1998; Sims e al. 1999). In Thailand,
11 species ha e been epo ed, namely S. a eola um, S. au an ium, S. bo is a,
S. cepa, S. ci inum, S. dic yospo um, S. la idum, S. lycope doides, S. poly hi-
zum, S. sinnama iense, and S. e ucosum, based on mo phological cha ac-
e is ics (Chand as ikul e al. 2011) and molecula analysis o S. su hepense
(Kumla e al. 2013).
In ou s udy, ou aim was o desc ibe h ee new species and a new eco d o
Scle ode ma om no he n Thailand, based on mac o- and mic oscopic cha -
ac e is ics and molecula phylogene ic me hods.
Ma e ials and me hods
Sample collec ion
F esh basidiomes o Scle ode ma we e collec ed du ing he ainy season
om May o June 2019 in Chiang Mai and Chiang Rai p o inces. A o al o 21
specimens we e collec ed, including ep esen a i es o h ee new species.
The specimens we e d ied in an o en a 45 °C o a leas 24 h and s o ed
a oom empe a u e. They a e deposi ed a Mae Fah Luang Uni e si y Fun-
ga ium (MFLU Funga ium).
Mo phological analysis
F esh basidioma a we e desc ibed o mac ocha ac e s, pho og aphed in he
ield, and es ed o mac ochemical eac ions (colo eac ions) o he pe idi-
um wi h 5% po assium hyd oxide (KOH). Thei size and colo we e eco ded,
and colo was compa ed wi h he Me huen Handbook o Colo (Ko ne up and
Wansche 1981). Fo obse a ion o mic ocha ac e s, d ied samples we e
sec ioned wi h azo blades, moun ed on glass slides, and ehyd a ed wi h
5% KOH (w/ ). The sizes and shapes o he mic os uc u es, including hy-
phae, basidia, and basidiospo es, we e examined unde a mic oscope (Nikon
DS-Ri2). A leas wo specimens o each species we e measu ed, and each
cha ac e is ic was measu ed in a leas 50 eplica es. Spo e o namen a ion
was also examined.
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MycoKeys 123: 69–88 (2025), DOI: 10.3897/mycokeys.123.160438
Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
DNA ex ac ion, PCR ampli ica ion, and sequencing
Genomic DNA was ex ac ed om 100 mg o he ui ing body using he Biospin
Fungus Genomic DNA Ex ac ion Ki (Bioe Technology, Hangzhou, China) ollow-
ing he manu ac u e ’s p o ocol. The polyme ase chain eac ion (PCR) was used
o ampli y he in e nal ansc ibed space (ITS), he 28S la ge subuni egion o i-
bosomal DNA (LSU), he second la ges subuni o RNA polyme ase II ( pb2), and
he ansla ion elonga ion ac o 1-alpha ( e 1-α). The p ime pai s used we e:
ITS1-F and ITS4 o ITS (Whi e e al. 1990; Ga des and B uns 1993), LR0R and LR5
o LSU (Vilgalys and Hes e 1990; Whi e e al. 1990), RPB2-6F and RPB2-7cR o
pb2 (Rehne and Buckley 2005), and EF1-983F and EF1-1567R o e 1-α (Rehne
and Buckley 2005). The PCR cycling condi ions o ITS, LSU, pb2, and e 1-α
we e: 3 min a 94 °C; 35 cycles o 30 s a 94 °C, 30 s a 52 °C, and 1 min a 72 °C;
ollowed by 10 min a 72 °C. Fo e 1-α, he p og am was: 5 min a 95 °C; 35 cy-
cles o 1 min a 94 °C, 2 min a 52 °C, and 1.5 min a 72 °C; ollowed by 10 min a
72 °C. Sequencing o PCR-ampli ied p oduc s in bo h di ec ions was pe o med
by Sangon Biological Enginee ing Technology and Se ices (Shanghai).
Alignmen and phylogene ic analysis
Phylogene ic analysis and sequence di e gence we e used o de e mine he
ela ionship o he newly disco e ed axon o o he Scle ode ma species. In he
phylogene ic analysis, Scle ode ma species om b oade geog aphic egions
we e included o compa ison. BioEdi Sequence Alignmen Edi o e sion
7.0.9.0 was used o e i y he ITS, LSU, pb2, and e 1-α sequences, and SeqMan
(DNAs a , Madison, WI, USA) was used o assemble he sequences. The da a-
base o he Na ional Cen e o Bio echnology In o ma ion (NCBI) (h p://www.
ncbi.nlm.nih.go /genbank/) was sea ched agains each sequence using he
Basic Local Alignmen Sea ch Tool (BLAST) o ensu e ha i belonged o he
co ec genus and was no con amina ed and o iden i y he closes ma ches.
A GenBank BLAST sea ch was pe o med o check o simila i y be ween he
newly gene a ed sequences. In o al, 82 sequences o se e al Scle ode ma spe-
cies om a ious egions, including ou collec ions, we e ob ained (Table 1).
Pisoli hus au an ioscab osus, a close ela i e o Scle ode ma (Ma in e al. 2002;
Wilson e al. 2012), was chosen as an ou g oup. Alignmen s we e pe o med
using MAFFT . 7.11 (h ps://ma .cb c.jp/alignmen /so wa e/, accessed on
23 Decembe 2023), and all alignmen s we e immed sepa a ely using T imAl
o elimina e ambiguously aligned posi ions (Capella-Gu ié ez e al. 2009).
The cha ac e se s included 78 collec ions and 676 cha ac e s (including
gaps) om ITS, 20 collec ions and 1402 cha ac e s om LSU, 20 collec ions
and 1120 cha ac e s om pb2, and 20 collec ions and 1035 cha ac e s om
e 1-α. The inal da ase comp ised 82 collec ions and 4233 cha ac e s om
ITS+LSU+ pb2+ e 1-α. A e checking o unsuppo ed con lic s (BS < 70%)
be ween single-gene maximum likelihood (ML) phylogenies, a conca ena ed
ou -locus da ase was assembled.
Phylogene ic analysis using ML was pe o med, ollowed by manual adjus -
men s in axmlGUI 2.0.13, along wi h Bayesian analysis, bo h conduc ed on he
CIPRES Science Ga eway e sion 3.3 web se e (Mille e al. 2010), a ailable a
h ps://www.phylo.o g/. A mixed-model (pa i ioned) scheme was employed o
72
MycoKeys 123: 69–88 (2025), DOI: 10.3897/mycokeys.123.160438
Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
bo h ML and Bayesian analyses, wi h he alignmen spli in o ou -cha ac e se s:
ITS, LSU, pb2, and e 1-α. The bes - i subs i u ion models o jModelTes 2 e -
sion 2.1.6 (Da iba e al. 2012) in XSEDE we e chosen o Bayesian analysis. The
selec ed models we e HKY+G o ITS, GTR+I+G o n LSU, HKY+G o pb2, and
SYM+I+G o e 1-α. Fou independen uns, each wi h ou chains, we e pe o med
o 1,000,000 gene a ions, wi h sampling e e y 100 gene a ions. The a e age s an-
da d de ia ion o spli equencies a he end o he uns was 0.015009. The bu n-in
phase (25%) was de e mined by assessing s a iona i y in he gene a ion-likelihood
plo using T ace e sion 1.7.1 (Rambau e al. 2018). The esul ing phylogene ic
ee was isualized in T ee iew 32 and u he edi ed using Adobe Illus a o CS6.0.
Table 1. Phylogene ic analysis lis o species, he ba ium numbe , place o o igin, and GenBank accession numbe .
Species Vouche in o ma ion Loca ion GenBank accession no. Re e ence
ITS LSU Rpb2Te 1-α
Pisoli hus
au an ioscab osus
AWW297 Malaysia EU718112 EU718146 FJ536648 FJ536681 Wilson e al. (2011)
Scle ode ma a eola um AWW211 USA EU718115 EU718149 FJ536651 FJ536683 Wilson e al. (2011)
S. a eola um PBM2208 Aus alia N/A EU718150 FJ536652 FJ536684 Wilson e al. (2011)
S. a eola um TNS:F-82295 Japan OQ025272 OQ025269 N/A N/A Kasuya e al. (2023)
S. a eola um Kasuya-B4422 Japan OQ025273 OQ025270 N/A N/A Kasuya e al. (2023)
S. a eola um O3C_4 USA JX030282 N/A N/A N/A Bzdyk e al. (2018)
S. a eola um 23 Spain MN684210 N/A N/A N/A -
S. a eola um Db-K -MH040288 N/A N/A N/A Bzdyk e al. (2018)
S. a eola um Bk-N -MH040301 N/A N/A N/A Bzdyk e al. (2018)
S. be mudense BZ3961 Belize EU718118 DQ644137 FJ536654 FJ536686 Wilson e al. (2011)
S. be mudense EUA09 -OQ351725 N/A N/A N/A Bullaín-Gala dis e al. (2024)
S. be mudense SUA03 -OQ351729 N/A N/A N/A Wilson e al. (2011)
S. bo is a MCA242 USA EU718117 DQ644138 FJ536653 FJ536685 Wilson e al. (2011)
S. ci inum AWW212 USA EU718119 EU718151 FJ536655 FJ536687 Wilson e al. (2011)
S. ci inum F-PRL5772 USA GQ166907 N/A N/A N/A Zhang e al. (2013)
S. ci inum K (M) 17485 England EU784413 N/A N/A N/A Zhang e al. (2013)
S. ci inum CITSCL1 USA FM213344 N/A N/A N/A Zhang e al. (2013)
S. ci inum K (M) 53906 England EU784414 N/A N/A N/A Zhang e al. (2013)
S. columna e CUB:Mic obiology KHS3 Thailand AB459512 N/A N/A N/A Ruankaew Disya a e al. (2016)
S. columna e Scl1 Thailand AB854700 N/A N/A N/A Kaewg ajang e al. (2023)
S. columna e CUB:Mic obiology KHS10 Thailand AB459519 N/A N/A N/A Ruankaew Disya a e al. (2016)
S. columna e MFLU25-0110 (DG150) Thailand N/A N/A PX137624 PX126632 This s udy
S. columna e MFLU25-0111 (DG153) Thailand PV444716 N/A PX137625 PX126633 This s udy
S. dic yospo um IR250 Bu kina Faso FJ840444 N/A N/A N/A Sanon e al. (2009)
S. dic yospo um IR408 Bu kina Faso FJ840445 N/A N/A N/A Sanon e al. (2009)
S. me idionale AWW218 USA EU718121 EU718152 FJ536656 FJ536688 Wilson e al. (2011)
S. mcalpinei OSC 24605 -EU718122 DQ682999 FJ536657 N/A Wilson e al. (2011)
S. ni idum UFRN:Fungos 2034 B azil KU759904 KU759903 N/A N/A Rau e al. (2020)
S. ni idum UFRN:Fungos 2219 B azil KU759908 N/A N/A N/A Rau e al. (2020)
S. poly hizum AWW216 USA EU718123 EU718153 FJ536658 FJ536689 Wilson e al. (2011)
S. poly hizum MA:Fungi-39352 Spain MT270662 N/A N/A N/A O iz-Ri e o e al. (2021)
S. sepa a um Ge5394 China OQ554975 N/A N/A N/A Wu e al. (2023)
S. sepa a um ZLR31 China OQ554974 N/A N/A N/A Wu e al. (2023)
S. sepa a um Ge4148 China OQ554973 N/A N/A N/A Wu e al. (2023)
S. sepa a um MFLU 19-1347 (NTF066) Thailand PV444715 PV446742 N/A N/A This s udy
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MycoKeys 123: 69–88 (2025), DOI: 10.3897/mycokeys.123.160438
Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
Species Vouche in o ma ion Loca ion GenBank accession no. Re e ence
ITS LSU Rpb2Te 1-α
S. sinnama iense SINSCL3 (SCLN) Thailand FM213358 N/A N/A N/A Phos i e al. (2009)
S. sinnama iense 150728-29 China MH513635 N/A N/A N/A Zhang e al. (2020)
S. sinnama iense SINSCL1 (SCLK4) Thailand FM213356 N/A N/A N/A Phos i e al. (2009)
S. sinnama iense SINSCL6 (SCLD1) Thailand FM213361 N/A N/A N/A Phos i e al. (2009)
S. sinnama iense SINSCL4 (SCLY5) Thailand FM213359 N/A N/A N/A Phos i e al. (2009)
S. sinnama iense CMU53:210-2 Thailand HQ687222 N/A N/A N/A Kumla e al. (2014)
S. sinnama iense p -355 -MW374160 N/A N/A N/A Wang e al. (2022)
S. sinnama iense HKAS122471 China ON794312 N/A N/A N/A Wang e al. (2022)
S. sinnama iense SINSCL5 (SC1) Thailand FM213360 N/A N/A N/A Phos i e al. (2009)
S. sinnama iense DX2021-8-2 -OL351633 N/A N/A N/A -
S. sinnama iense MFLU25-0112 (DG157) Thailand N/A N/A PX137633 PX207694 This s udy
S. sinnama iense MFLU25-0113 (DG158) Thailand PV444717 N/A PX137632 PX207695 This s udy
S. sinnama iense MFLU25-0114 (DG159) Thailand PV444718 N/A PX137631 PX137634 This s udy
S. sinnama iense MFLU25-0115 (DG160) Thailand PV444719 N/A PX137630 PX126634 This s udy
S. sinnama iense MFLU 19-1647 (MO-DG020) Thailand PV444720 N/A PX137626 PX126635 This s udy
S. sinnama iense MFLU 19-1648 (MO-DG021) Thailand PV444721 N/A N/A N/A This s udy
S. sinnama iense MFLU 19-1649 (MO-DG022) Thailand PV444722 PV446743 PX137627 PX126636 This s udy
S. sinnama iense MFLU 19-1650 (MO-DG023) Thailand PV444723 N/A PX137628 N/A This s udy
S. sinnama iense MFLU 19-1652 (MO-DG034) Thailand PV444724 N/A N/A N/A This s udy
S. sinnama iense MFLU 19-1653 (MO-DG035) Thailand PV444725 N/A PX137629 N/A This s udy
S. sinnama iense MFLU 19-1341 (NTF012) Thailand PV444726 N/A N/A N/A This s udy
Scle ode ma sp. AWW260 Malaysia EU718124 EU718155 FJ536660 FJ536691 Wilson e al. (2011)
Scle ode ma sp. AB96 Came oon KR819100 N/A N/A N/A Michaëlla Ebenye e al. (2017)
Scle ode ma sp. YAAS-L5455 -MT876542 N/A N/A N/A -
Scle ode ma sp. YAAS-L5449 -MT876541 N/A N/A N/A -
Scle ode ma sp. SL2085 Singapo e OR354966 N/A N/A N/A -
Scle ode ma sp. ECM26-SERS -DQ146385 N/A N/A N/A Yuwa-Amo npi ak e al. (2006)
Scle ode ma sp. LH35 Malaysia GQ268582 N/A N/A N/A Peay e al. (2010)
Scle ode ma sp. MFLU 19-1348 (NTF090) Thailand PV444728 N/A N/A N/A This s udy
Scle ode ma sp. MFLU 19-1517 (DMSL-
DG005)
Thailand PV444729 N/A N/A PV749898 This s udy
S. su hepense AWW311 Malaysia EU718125 EU718156 FJ536661 FJ536692 Wilson e al. (2011)
S. su hepense CMU:55-SC2 Thailand NR_132871 N/A N/A N/A Kumla e al. (2013)
S. su hepense JH-2016-0727-052 China MH513626 N/A N/A N/A Zhang e al. (2020)
S. su hepense 180508-08 China MH513625 N/A N/A N/A Zhang e al. (2020)
S. su hepense MFLU25-0109 (DG146) Thailand N/A N/A N/A PX126630 This s udy
S. su hepense MFLU 19-1344 (NTF053) Thailand PV444727 N/A N/A N/A This s udy
S. xan hoch oum AWW254 Malaysia EU718126 EU718154 N/A N/A Wilson e al. (2011)
S. yunnanense HKAS80386 -MW493647 MW493703 N/A N/A Kasuya e al. (2023)
S. yunnanense PERTH-7604645 China MT270651 N/A N/A N/A O iz-Ri e o e al. (2021)
S. yunnanense TNS:F-82294 Japan OQ025271 OQ025268 N/A N/A Kasuya e al. (2023)
uncul u ed ungus ASV_419 -LR993736 N/A N/A N/A -
uncul u ed ungus ASV_1014 -LR994331 N/A N/A N/A -
Scle ode ma longis ipes MFLU 19-1655 (DG109) Thailand PV444712 PV446740 PX126607 PX121227 This s udy
Scle ode ma longis ipes MFLU 19-1656 (DG110) Thailand PV444713 PV446741 PX121228 N/A This s udy
Scle ode ma
magnispo um
MFLU 19-1345 (NTF062) Thailand PV444714 N/A N/A N/A This s udy
Scle ode ma
mic oca pum
MFLU 19-1349 (DG002) Thailand PV436898 N/A N/A N/A This s udy
N/A: no a ailable; species in his s udy a e indica ed in bold black.
74
MycoKeys 123: 69–88 (2025), DOI: 10.3897/mycokeys.123.160438
Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
Resul s and discussion
Based on Scle ode ma species, hey we e o me ly sepa a ed by mo phology, wi h
basidiome size and shape a ying depending on soil and en i onmen and basid-
iospo e mo phology (Wa ling 2006; Sanon e al. 2009; Kumla e al. 2013; Gonk-
hom e al. 2025). In his s udy, Scle ode ma sp. no ., desc ibed om mac omo -
phological and mic omo phological ai s oge he wi h phylogene ic analysis o
ITS, LSU, pb2, and e 1-α genes, includes h ee new species—Scle ode ma longis-
ipes, Scle ode ma magnispo um, and Scle ode ma mic oca pum— om no he n
Thailand. This wo k enhances ou unde s anding o he di e si y o Scle ode ma
species. Mac oscopically, he size, colo , and ype o dehiscence o he basidi-
ome, as well as he colo and hickness o he pe idium, a e c ucial ai s o iden i-
ying Scle ode ma. Mic oscopically, he size, shape, and o namen a ion o he ba-
sidiospo es a e employed o di e en ia e species o his genus. The basidiomes
and basidiospo es a e simila in size and cha ac e o hose o S. sepa a um,
S. dic yospo um, and S. hypogaeum (Sims e al. 1995; Co ez e al. 2011; Guzmán
e al. 2013). Ou ITS, LSU, pb2, and e 1-α sequence analyses clea ly sepa a e
hese species om o he e icula e-spo ed Scle ode ma species in he sec ion.
Phylogene ic analysis
The combined da ase o ou genes comp ised 4233 bp (including gaps):
676 bp o ITS, 1402 bp o LSU, 1120 bp o pb2, and 1035 bp o e 1-α. The
bes RAxML phylog am, wi h a inal likelihood alue o -22414.937168, is p e-
sen ed. The ma ix had 1563 dis inc alignmen pa e ns wi h 68.28% unde e -
mined cha ac e s o gaps. The es ima ed base equencies we e: A = 0.233328,
C = 0.255317, G = 0.274489, and T = 0.236866. Subs i u ion a es we e AC =
1.330307, AG = 4.855565, AT = 1.388503, CG = 1.501262, CT = 8.964382, and
GT = 1.000000. The gamma dis ibu ion shape pa ame e was α = 0.929050.
The phylog am opology de i ed om Bayesian analysis was simila o ha
om ML analysis. Boo s ap alues o ML ≥ 70% and Bayesian pos e io p oba-
bili ies (PP) ≥ 0.90 a e indica ed in Fig. 1.
Phylogene ic ees in e ed om ML and M Bayes analyses esul ed in sim-
ila opologies; he e o e, only he ML ee (ITS+LSU+ pb2+ e 1-α) is shown,
wi h bo h ML boo s ap (BS) alues and Bayesian pos e io p obabili ies (PP).
In he phylog am, Scle ode ma longis ipes (MFLU 19-1655 and MFLU 19-1655)
was closely ela ed o Scle ode ma sp. (uncul u ed ec omyco hiza ECM26
SERS, LH35 om Malaysia, and SL2085 om Singapo e) wi h high s a is ical
suppo : 100% BS/1.00 PP, 81% BS/0.93 PP, and 90% BS/1.00 PP, espec i ely.
Scle ode ma mic oca pum (MFLU 19-1347) was closely ela ed o he uncul-
u ed ungi ASV 1014 and ASV 419, wi h s a is ical suppo o 70% BS/0.94 PP.
Scle ode ma magnispo um (MFLU 19-1345) was closely ela ed o Scle ode ma
sp. YAAS L5449 and YAAS L5455, wi h s a is ical suppo o 74% BS/0.97 PP.
The species mos closely ela ed in he phylogene ic ee o Scle ode ma
mic oca pum (MFLU 19-1347) was S. dic yospo um (Vouche IR250), wi h a
gene ic dis ance o 11.81% (65/570) be ween ITS sequences. Scle ode ma
magnispo um (MFLU 19-1345) and S. mic oca pum (MFLU 19-1347) showed a
gene ic dis ance o 21.34% (95/445), suppo ing hei dis inc ion as sepa a e
species. This esul is consis en wi h p e ious molecula phylogene ic s udies
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Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
Figu e 1. Phylogene ic ee ob ained om he maximum likelihood analysis o Scle ode ma species. Maximum likelihood
ee ob ained om he alignmen o ITS, LSU, pb2, and e 1-α sequences. The boo s ap consensus ee was in e ed
om 1000 eplica es. Pisoli hus au an ioscab osus was included as an ou g oup.
ha s ongly suppo he ecogni ion o Scle ode ma species as gene ically dis-
c e e lineages (Phos i e al. 2009; Nouh a e al. 2012; Wu e al. 2023).
Scle ode ma species a e ound in empe a e, opical, and sub opical egions,
which may be ela ed o he highe di e si y o Scle ode ma o ec omyco hizal
ungi in hese clima ic zones (Je ies 1999; B und e e al. 2005; P adhan e
al. 2011; Wu e al. 2023). In addi ion, 11 species o Scle ode ma (S. a eola um,
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Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
S. bo is a, S. cepa, S. ci inum, S. dic yospo um, S. la idum, S. lycope doides, S.
poly hizum, S. sinnama iense, S. e ucosum, and S. su hepense) ha e been e-
co ded in Thailand based on mo phology (Chand as ikul e al. 2011; Kumla e al.
2013; Gonkhom e al. 2025). Phylogene ic analysis con i med he placemen o
Scle ode ma sepa a um based on he ITS and LSU egions (Fig. 1). S. sepa a um
exhibi s simila shapes and sizes o basidiomes and basidiospo es (Wu e al.
2023). Phylogene ic analysis based on ITS and LSU sequences acili a ed con i -
ma ion o he species we analyzed, which has been o icially eco ded in Thailand.
Taxonomy
Scle ode ma longis ipes Gonkhom, Sysouph. & Thongkl., sp. no .
Index Fungo um: IF903880
Fig. 2
Diagnosis. Epigeous b own o bu n umbe basidioma a wi h long s ipe, ub-
be y pale b own pe idium, hyaline o yellow b own hyphae in exope idium, hy-
aline hyphae in endope idium, globose da k b own basidiospo es wi h echinu-
la e o spinose o namen a ion.
Holo ype. Thailand • Chiang Rai P o ince, Mueang Chiang Rai Dis ic , Mae
Fah Luang Uni e si y campus, 04 June 2019, collec ed by Didsanu da Gonk-
hom, DG109 (MFLU 19-1655).
E ymology. The species name (longis ipes) e e s o he long s ipe o he
basidioma a.
Desc ip ion. Basidioma a epigeous, 28–35 mm in diam., 42–60 mm high,
club-shaped, wi h globula pe idial head; wi h c acked o squamulose su ace,
b own (6E5) backg ound when young, wi h awn (7E4) o b own (6E5) o bu n
umbe (6F6) squamula c acks upon lu eous backg ound, ha d skin, ough when
ma u e. S ipe sub-cylind ic, a , wi h small i egula c acks a he op o he
s ip, 35–45 × 10–13 mm, whi e backg ound, co e ed wi h b own (6E5) ib il-
lose squamules. Rhizomo phs mo e agg ega ed a he base, whi e, b anched,
and na owing owa ds he base. Con ex whi e in pe idium and s ipe, u ned
dull ed o g eyish ed (98B4-5) when cu . Pe idium up o 5 mm wide when
esh, ubbe y in consis ence, pale b own (6D5).
Pe idium laye o med by simple-sep a e hyphae. Exope dium sligh ly hick-
ened walls, composed o in e wo en o ami ied and supe imposed hyphae,
hyaline o yellow b own, 2.9–3.8 µm diam. Endope dium hick, composed o in-
e wo en hyphae, hyaline, 4.3–7.2 µm diam. Clamp connec ions p esen on endo-
pe idium hyphae. Gleba b ownish g ey (9E2), g eyish b own (9E3), o oxblood ed
(9E7) o da k b own (9F4-7), compac , and powde y when ma u e. Basidiospo es
(n = 50) globose, echinula e, da k b own in KOH, (13.4–)14.5–17.2(19.5) µm in
diam., wi h b own spinose o namen a ion (2.6–5.9 µm high). Basidia no seen.
Habi a and dis ibu ion. Caespi ose o ascicula ed on soil, epigeous, in
no he n Thailand.
Addi ional specimens examined. Thailand • Chiang Rai P o ince, Mueang
Chiang Rai Dis ic , Mae Fah Luang Uni e si y campus, 04 June 2019, collec ed
by Didsanu da Gonkhom, DG110 (MFLU 19-1656).
No es. Scle ode ma longis ipes is cha ac e ized by a la ge b own basidioma-
a wi h a longe s ipe ha u ns om dull ed o g eyish ed when ouched. The
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Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
Figu e 2. Scle ode ma longis ipes (MFLU 19-1655, holo ype). A, B. Basidioma a; C. Scale on pe idium su ace; D. Cu
side o pe idium o MFLU 19-1655; E, F. Basidioma a; G. Scale on pe idium su ace; H. Cu side o pe idium o Scle o-
de ma longis ipes (MFLU 19-1656); I. Exope idial hyphae; J. Endope idial hyphae; K. Clamped hyphae o endope idium;
L–Q. Basidiospo e. Scale ba s: 10 mm (A, B, E, F); 5 mm (C, D, G, H); 50 µm (I, J); 20 µm (K); 10 µm (L–Q).
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Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
3 Pe idium yellowish o o angish-yellow, smoo h o c acked ..............S. cepa
– Pe idium b own, wi h dis inc wa s o scales ............................................4
4 Pe idium co e ed wi h ough aised wa s, yellow-b own ..........S. ci inum
– Pe idium c acked, scaly, oughened, b own.................................. S. bo is a
5 Pe idium hin, lea he y, yellowish whi e .......................................................6
– Pe idium hick, wi h dis inc c acks o subscaly .........................................8
6 Basidiospo es 7–8.5 µm, da k b own, e icula e ................S. dic yospo um
– Basidiospo es la ge han 8.5 µm, spiny ......................................................7
7 Basidiospo es 8–12 µm, ound, ne -like idges .....................S. e ucosum
– Basidiospo es 10–15 µm, sligh ly oughened ex u e ............S. columna e
8 Pe idium smoo h, b own, ough, hick (up o 5 mm) ......... S. magnispo um
– Pe idium c acked o scaly, backg ound yellowish.......................................9
9 Basidiospo es 8.24–13.33 µm, hizomo phs pale b own .....S. mic oca pum
– Basidiospo es 12.31–16.49 µm, spines up o 3.9 µm ............S. sepa a um
10 S ipe well-de eloped, mo e han 3 cm long ...............................................11
– S ipe sho , less han 3 cm, o absen ........................................................13
11 S ipe sub-cylind ic, c acked a he op, basidiospo es 13.4–19.5 µm .........
...................................................................................................S. longis ipes
– S ipe sho o i egula , basidiospo es smalle (<14 µm) .........................12
12 Basidiospo es 7–12 µm, pe idium s a -shaped when spli .......S. poly hizum
– Basidiospo es 8–13 µm, pe idium smoo h o scaly ............. S. su hepense
13 Pe idium golden yellow, apex up u ing a ma u i y ................... S. la idum
– Pe idium b own o och aceous, scaly, o oughened ................................14
14 Pe idium lea he y, e ucose, yellowish o lemon-yellow ......S. sinnama iense
– Pe idium b ownish, smoo h o scaly, spo e mass da k b own .................15
15 Basidiome su ace smoo h, spo e mass clea ly oli e ......S. lycope doides
– Basidiome ough o scaly, spo e mass b own o da k b own ..... S. a eola um
O e all, his s udy signi ican ly ad ances ou unde s anding o he di e si y
and axonomy o Scle ode ma species in no he n Thailand, desc ibing h ee
new species—Scle ode ma longis ipes, S. mic oca pum, and S. magnispo um—
and epo ing a new eco d, S. sepa a um, om Thailand. The esea ch in e-
g a es comp ehensi e mo phological analyses wi h molecula phylogene ic
me hods, u ilizing sequences om ou loci (ITS, LSU, pb2, and e 1-α) o con-
i m he dis inc i eness o hese axa. The indings con ibu e o ou knowledge
o Scle ode ma and es ablish a basis o u u e ecological and e olu iona y e-
sea ch on ec omyco hizal ungi.
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
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Didsanu da Gonkhom e al.: Th ee new Scle ode ma species and a eco d om no he n Thailand
Funding
The au ho s app ecia e he kind suppo p o ided by he Resea ch and Resea che s
Indus ies (RRI) (g an no. PHD62I0018/2562) and Mae Fah Luang Uni e si y.
Au ho con ibu ions
Didsanu da Gonkhom: me hodology, da a cu a ion, w i ing—o iginal d a . Phongeun
Sysouphan hong: in es iga ion, concep ualiza ion, w i ing. Ma c S adle , Na i sada
Thongklang, and Ke in D. Hyde: w i ing— e iew and edi ing, supe ision. All o he au-
ho s ead, p o ided use ul edi s, and ag eed o he published e sion o he manusc ip .
Au ho ORCIDs
Didsanu da Gonkhom h ps://o cid.o g/0000-0002-4827-9771
Phongeun Sysouphan hong h ps://o cid.o g/0000-0002-1275-2459
Ma c S adle h ps://o cid.o g/0000-0002-7284-8671
Na i sada Thongklang h ps://o cid.o g/0000-0001-9337-5001
Ke in D. Hyde h ps://o cid.o g/0000-0002-2191-0762
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex .
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