147
Th ee new species o Fuscopo ia (Hymenochae ales,
Basidiomyco a) om sou he n India e ealed by mo phological
and mul igene phylogene ic analyses
Sugan ha Gunaseelan1, Kezhocuyi Kezo1, Elango an A umugam1, Saman ha C. Ka una a hna2,
Al-Banda i Fahad Al-A jani3, Abdallah M. Elgo ban4, Ja u ong Kumla5, Naka in Suwanna ach5,
Ekachai Chukea i o e6, Mala izhi Kaliyape umal1
1 Cen e o Ad anced S udies in Bo any, Uni e si y o Mad as, Guindy Campus, Chennai 600025, Tamil Nadu, India
2 Cen e o Yunnan Pla eau Biological Resou ces P o ec ion and U iliza ion & Yunnan In e na ional Join Labo a o y o Fungal Sus ainable U iliza ion in Sou h
and Sou heas Asia, College o Biology and Food Enginee ing, Qujing No mal Uni e si y, Qujing 655099, China
3 Bo any and Mic obiology Depa men , College o Science, King Saud Uni e si y, Riyadh 11451, Saudi A abia
4 Cen e o Excellence in Bio echnology Resea ch (CEBR), DSR, King Saud Uni e si y, Riyadh, Saudi A abia
5 Cen e o Excellence in Mic obial Di e si y and Sus ainable U iliza ion, Chiang Mai Uni e si y, Chiang Mai 50200, Thailand
6 School o Science, Mae Fah Luang Uni e si y, Chiang Rai, 57100, Thailand
Co esponding au ho s: Ekachai Chukea i o e ([email p o ec ed]); Mala izhi Kaliyape umal (mala [email p o ec ed])
Copy igh : © Sugan ha Gunaseelan 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
Fuscopo ia Mu ill, a cosmopoli an genus o he Hymenochae aceae, consis s o pa -
asi ic and sap o ophic ungi cha ac e ized by esupina e o pilea e, s ic ly dimi ic hy-
phal sys ems, enc us ed gene a i e hyphae, he p esence o hymenial se ae, and hyaline,
smoo h, hin-walled basidiospo es. Based on mo pho-mic o axonomic examina ions
and phylogene ic analyses using a combined ITS, n LSU, pa ial pb2, and e 1-α da ase ,
h ee new species o Fuscopo ia—F. indica, F. si umalaiensis, and F. e minalianae—a e de-
sc ibed om he Eas e n Gha s o Tamil Nadu, India. The newly desc ibed species o m
h ee dis inc lineages wi hin he F. o ulosa complex. Fuscopo ia indica is cha ac e ized
by imb ica e, con ex, dimidia e basidiomes, a glab ous, azona e pilea su ace, and el-
lipsoid basidiospo es (3.5–4.5 × 2.5–2.9 μm). Fuscopo ia si umalaiensis has a smoo h
o glab ous, concen ically zona e pilea su ace, a duplex con ex , and smalle basidio-
spo es (3.1–3.6 × 2.1–2.6 μm). Finally, F. e minalianae is ecognized by i s e used- e-
lexed o imb ica e basidiome, widely zona e and wa ed pilea su ace, and ellipsoidal
basidiospo es (3.3–4.3 × 2.8–3 μm). This s udy p o ides comp ehensi e desc ip ions,
mo phological illus a ions, and insigh s in o he di e ences among hese new species
and hei espec i e allied axa, along wi h he esul s o phylogene ic analysis.
Key wo ds: Hymenochae aceae, mul igene phylogeny, Phellinus sensu la o, axonomy,
wood- o ing ungi
In oduc ion
The genus Fuscopo ia Mu ill was e ec ed wi h F. e uginosa (Sch ad.) Mu ill
as he ype species (Mu ill 1907). I is widely ecognized as a o es pa hogen
ha causes whi e o in bo h coni e ous and deciduous ees (Panconesi e
Academic edi o : R. Hen ik Nilsson
Recei ed:
8 Augus 2025
Accep ed:
20 Oc obe 2025
Published:
18 No embe 2025
Ci a ion: Gunaseelan S, Kezo K,
A umugam E, Ka una a hna SC, Fahad
Al-A jani A-B, Elgo ban AM, Kumla
J, Suwanna ach N, Chukea i o e
E, Kaliyape umal M (2025) Th ee
new species o Fuscopo ia
(Hymenochae ales, Basidiomyco a)
om sou he n India e ealed by
mo phological and mul igene
phylogene ic analyses. MycoKeys 125:
147–166. h ps://doi.o g/10.3897/
mycokeys.125.168173
MycoKeys 125: 147–166 (2025)
DOI: 10.3897/mycokeys.125.168173
148
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
al. 1994; Spi in e al. 2014; Luana e al. 2015). Fuscopo ia is cha ac e ized by
i s annual o pe ennial, esupina e o pilea e basidiomes, a dimi ic hyphal sys-
em wi h enc us ed gene a i e hyphae, and he p esence o hymenial se ae in
mos species excep o F. sho eae, F. longise ulosa, and F. discipes. Ma ginal
se ae may be p esen o absen , and he basidiospo es a e hyaline, hin-walled,
smoo h, and ange in shape om subglobose o cylind ical (Dai 2010; Wu e
al. 2022). Fuscopo ia is dis ibu ed wo ldwide; mo e han 1,561 nucleo ide se-
quences belonging o 63 known and 55 unspeci ied species a e publicly a ail-
able in GenBank (Saye s e al. 2025), and 130 axa a e egis e ed in MycoBank
(Robe e al. 2013) as o Oc obe 2025.
Phylogene ic s udies o No h Ame ican and Eu opean membe s o he Hy-
menochae aceae based on n LSU ede ined Phellinus sensu la o in o se e al
monophyle ic gene a. These include Fomi ipo ia Mu ill, Fomi ipo ella Mu ill,
Ful i omes Mu ill, Fuscopo ia Mu ill, Phellinus s. s ., Po odaedalea Mu ill,
Phellinidium (Ko l.) Fiasson and Niemelä, Phellopilus Niemelä, T. Wagne and
M. Fisch., Au i ica ia D.A. Reid, Onnia P. Ka s ., and Phyllopo ia Mu ill (Fias-
son and Niemelä 1984; Fische 1996; Niemelä e al. 2001; Wagne and Fische
2001, 2002), all o which we e la e accep ed as dis inc axa (Niemelä e al.
2001; Spi in e al. 2006; G oposo e al. 2007; Bal aza e al. 2009; Bal aza and
Gibe oni 2010; Dai 2010; Raymundo e al. 2013; Chen and Yuan 2017; Chen e
al. 2019; Chen and Dai 2019). Recen molecula analyses o Fuscopo ia ha e
e ealed ha wi hin his axon, he e a e se e al majo g oupings suppo ed by
subs an ial mo phological and phylogene ic e idence. These a e he F. con i-
gua complex, he F. e a complex, he F. e uginosa complex, he F. gil a com-
plex, he F. o ulosa complex, and he F. i icola complex (Chen e al. 2022, 2024;
Wu e al. 2022; Cho e al. 2023).
Chen e al. (2019, 2020, 2023a, b) p oposed 18 new species—F. acu ima gin-
a a, F. aus alasica, F. aus aliana, F. ame icana, F. bambusae, F. cen oame icana,
F. chinensis, F. cos a icana, F. eucalyp i, F. ka s eniana, F. la ispo a, F. mon icola, F.
plume iae, F. amulicola, F. sep ise a, F. sinica, F. sho eae, and F. subch ysea— om
Aus alia, China, Cos a Rica, Mexico, Singapo e, Thailand, and he Uni ed S a es.
F om 2022 o 2024, nume ous new species—F. dolichose a, F. dollinge i, F. euca-
lyp icola, F. gil oides, F. hawaiiana, F. ko eana, F. minu issima, F. nadi i ana, F. esupi-
na e, F. e icula a, F. semicephala, F. sinuosa, F. submu ina, and F. sub opica—ha e
been epo ed wo ldwide (Wu e al. 2022; Chen e al. 2023a, b; Cho e al. 2023;
Chen e al. 2024; C ous e al. 2024). Recen ly, Chen e al. (2025) epo ed Fuscopo-
ia e lexoides, a new species belonging o he F. gil a complex om China.
Despi e signi ican ad ancemen s, se e al egions o he wo ld emain un-
de -sampled, including Oceania, pa s o Sou h Ame ica, opical A ica, and
Sou heas Asian islands. Fo a mo e comp ehensi e unde s anding o Fusco-
po ia, widesp ead sampling om Neo opical, Paleo opical, opical, and em-
pe a e coun ies o Asia is essen ial. Ea lie , knowledge o he species di e -
si y o hymenochae oid ungi in India was con ined o he no he n egions,
p ima ily based on mo pho- axonomic cha ac e is ics. Kuma i e al. (2021)
documen ed 14 species o Phellinus (P. callimo phus, P. ch yseus, P. con iguus,
P. discipes, P. e eus, P. e uginosus, P. gil us, P. o ien alis, P. punc a i o mis,
P. haba ba inus, P. u i inc us, P. senex, P. o ulosus, and P. wahlbe gii), which
ha e since been alida ed and ans e ed o Fuscopo ia h ough phylogene ic
analysis. F om he sou he n egion o India, Bakshi (1971) epo ed P. discipes
149
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
(syn. F. discipes). Ou p e ious s udy ep esen ed he i s mul igene app oach
in e ealing F. nadi i ana om he Eas e n Gha s o Tamil Nadu (C ous e al.
2024). In he p esen s udy, we epo h ee no el species o Fuscopo ia om
he Eas e n Gha s o Tamil Nadu and p o ide de ailed desc ip ions, illus a ions,
and phylogene ic placemen based on ITS+n LSU+ pb2+ e 1-α sequence da a.
Ma e ials and me hods
Mo phological analyses
The s udied specimens we e collec ed om a ious loca ions in he Eas e n
Gha s o Tamil Nadu, India. Mo phological and mic oscopic cha ac e is ics
we e analyzed as desc ibed ea lie (Gunaseelan e al. 2024), and he phene -
ic colo codes ollow he Me huen Handbook (Ko ne up and Wansche 1978).
F eehand sec ions o d ied basidiomes we e moun ed in wa e , 5% po assium
hyd oxide (KOH), co on blue (CB), and Melze ’s eagen (IKI) o analyze mi-
c o-mo phological ea u es. Mic oscopic obse a ions, measu emen s, and
line d awings we e pe o med in 5% KOH using a LABOMED CxL2 compound
mic oscope. Pho omic og aphs we e aken wi h a LABOMED OPTIC-CX BINO
LED mic oscope a magni ica ions up o 1000×. Basidiospo e measu emen s
(minimum–mean–maximum) and Q alues (leng h/wid h a ios) we e eco d-
ed. Abb e ia ions: CB− = acyanophilous, CB+ = cyanophilous, IKI− = inamyloid,
IKI+ = amyloid, Q = L/W a io (basidium leng h excluding s e igma a), L = mean
spo e leng h (a i hme ic mean o all spo es), W = mean spo e wid h (a i hme-
ic mean o all spo es), n = numbe o spo es measu ed. Measu emen s we e
based on 50 basidiospo es, 30 cys idioles, hymenial se ae, basidioles, and ba-
sidia pe specimen. The iden i ied specimens we e deposi ed a he Mad as
Uni e si y Bo any Labo a o y (MUBL), Cen e o Ad anced S udies in Bo any,
Uni e si y o Mad as, Chennai 600025, Tamil Nadu, India.
Genomic DNA ex ac ion, PCR ampli ica ion, and sequencing
The nuclea ibosomal in e nal ansc ibed space (ITS) egion was ampli ied
using p ime s ITS1 and ITS4 (Whi e e al. 1990). PCR ampli ica ion was ca ied
ou unde he ollowing condi ions: ini ial dena u a ion a 95 °C o 3 minu es,
ollowed by 32 cycles o 95 °C o 30 seconds, 52 °C o 30 seconds, and 68 °C
o 1 minu e, wi h a inal ex ension a 68 °C o 3 minu es. The nuclea ibosom-
al la ge subuni (n LSU) egion was ampli ied using p ime s LR0R and LR7 (Vilg-
alys and Hes e 1990). PCR ampli ica ion condi ions o n LSU we e as ollows:
ini ial dena u a ion a 94 °C o 1 minu e, ollowed by 34 cycles a 94 °C o 30
seconds, 45 °C o 30 seconds, and 72 °C o 1.5 minu es, and a inal ex en-
sion a 72 °C o 10 minu es. The RNA polyme ase II gene ( pb2) was ampli ied
wi h p ime s bRPB2-6F and bRPB2-7.1R (Ma heny 2005), and PCR condi ions
we e as ollows: ini ial dena u a ion a 94 °C o 2 minu es; 35 cycles o 94 °C
o 45 seconds, 53 °C o 90 seconds, and 72 °C o 90 seconds; and a inal
ex ension a 72 °C o 10 minu es. The ansla ion elonga ion ac o 1-alpha
gene ( e 1-α) was ampli ied wi h p ime pai EF1-983F/EF1-1567R (Rehne and
Buckley 2005). PCR ampli ica ion condi ions o e 1-α we e as ollows: ini ial
dena u a ion a 94 °C o 3 minu es, ollowed by 34 cycles o dena u a ion a
150
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
94 °C o 40 seconds, annealing a 54 °C o 45 seconds, ex ension a 72 °C o
1 minu e, and a inal ex ension a 72 °C o 10 minu es. Sequencing was ou -
sou ced o Eu o ins Genomics India P . L d., Ka na aka, India.
Phylogene ic analyses
Fo he phylogene ic analyses, conca ena ed sequences (ITS+n LSU+ p-
b2+ e 1-α) gene a ed om his s udy and allied axa e ie ed om GenBank
(Table 1), along wi h he ou g oups Coni e ipo ia wei ii (CFS 504) and Phellinidi-
um ag ans (CBS 202.90), we e aligned in MEGA X 10.0.2, ollowed by manual
adjus men s o enhance alignmen accu acy (Kuma e al. 2018). To acili a e
da ase pa i ioning, he mos app op ia e pa i ioning scheme and subs i u ion
models we e de e mined using Pa i ionFinde 1.10 (Lan ea e al. 2012) unde
he AIC c i e ion wi h he “g eedy” sea ch op ion. The da ase was subdi ided
in o 10 pa i ions: ITS1, 5.8S, ITS2, n LSU, pb2 in ons, pb2 exon, e 1-α in ons,
e 1-α 1s codon, e 1-α 2nd codon, and e 1-α 3 d codon. The bes - i e olu ion-
a y models selec ed by M Model es 2.3 (Nylande 2004) we e implemen ed
o each pa i ion: GTR+G (ITS1), GTR+I+G (5.8S), JC69 (ITS2), HKY85 (n LSU),
GTR+G ( pb2 in ons), GTR+G ( pb2 exon), HKY85+I+G ( e 1-α in ons), GTR+G
( e 1-α 1s codon), HKY85+G ( e 1-α 2nd codon), and K80+G ( e 1-α 3 d codon). The
combined da ase wi h pa i ion-speci ic models was used o Bayesian analysis
using M Bayes e sion 3.2.7a (Ronquis e al. 2012) wi h wo independen uns
comp ising ou Me opolis-coupled Ma ko chain Mon e Ca lo chains, each
unning o 4,000,000 gene a ions wi h sampling e e y 1,000 gene a ions. The
i s 25% o sampled ees we e disca ded as bu n-in, and he emaining ees
we e used o in e a majo i y- ule consensus and calcula e Bayesian pos e io
p obabili ies (BPP) o he clades. Maximum likelihood (ML) analyses we e pe -
o med using axmlGUI 2.0 (Edle e al. 2020), in ol ing 1,000 eplica es unde
he GTR+GAMMAI model. Addi ionally, 1,000 apid boo s ap eplica es we e
pe o med using he GTRCAT model o e alua e he ML boo s ap alues o he
nodes. A clade was conside ed s ongly suppo ed i boo s ap (BS) ≥ 60% and
pos e io p obabili y (PP) ≥ 0.80. Newly gene a ed sequences we e deposi ed in
GenBank (h p://www.ncbi.nlm.nih.go /) (Table 1). The alignmen was submi -
ed o Figsha e (DOI: 10.6084/m9. igsha e.29844794).
Resul s
Molecula phylogeny
In o al, 22 new sequences o ITS, n LSU, pb2, and e 1-α gene a ed in his
s udy we e submi ed o GenBank, and he accession numbe s a e lis ed
in Table 1. In addi ion, ITS, n LSU, pb2, and e 1-α sequences o 74 allied
axa (65 ITS, 72 n LSU, 28 pb2, and 39 e 1-α) we e e ie ed om Gen-
Bank (Table 1), along wi h he ou g oups Coni e ipo ia wei ii (CFS504) and
Phellinidium ag ans (CBS 202.90). The conca ena ed mul iple sequence
alignmen was 3,431 bases long, o which 1,854 we e cons an , 1,468 we e
a iable, and 1,130 (33%) we e pa simony in o ma i e. The ITS egion com-
p ised 879 bases, n LSU 1,385 bases, pb2 620 bases, and e 1-α 544 bases.
Maximum likelihood (ML) and Bayesian in e ence (BI) analyses gene a ed
151
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Table 1. Names, s ain numbe s, coun ies o collec ion, and co esponding GenBank accession numbe s o he se-
quences used in his s udy. Newly gene a ed sequences a e in bold, ‘T’ indica es ype specimens, and em dashes (—)
indica e da a una ailabili y.
Species S ain Coun y ITS LSU e 1-α pb2
Fuscopo ia acu ima gina aTDai 16892 China MH050752 MH050766 MN848822 MH079393
F. ambiguaTJV 0509/151 Uni ed S a es MN816707 MN809996 —MN848792
F. ame icanaTBJFC 020644 Uni ed S a es NR_173723 ———
F. a lan ica VRTO24 B azil ON808610 ON795835 — —
F. aus alasicaTDai 15636 China MG008397 MG008450 MH636408 MH079402
F. aus alianaTDai 18879 Aus alia MN816705 MN810015 MN848850 MN848767
F. bambusaeTDai 16599 Thailand MN816711 NG_075315 MN848808 —
F. bambusicolaTCui 8692 China MN816739 MT032486 MN848813 —
F. callimo pha Doll 868 China MN816701 MN809992 MN848840 —
F. caymanensisTJV 1908/74 F ench Guiana MT676832 MT676833 — —
F. cen oame icana JV 1606/93 Cos a Rica MG008444 MG008460 MH636389 —
F. chinensisTDai 15713 China MN816721 MN810008 MN848846 MN848771
F. ch ysea JV 1607/106-J Cos a Rica MN816736 MN810027 MN848818 MN848773
F. cinchonensis CBS 447.76 Sou h Ko ea AY558613 ———
F. con iguaTDai 16025 Uni ed S a es MG008401 MG008454 MH636386 MH079406
F. cos a icana JV 1407/92 Cos a Rica MG008446 MG008461 — —
F. dho a ensis ATN-007 Oman OP593104 OP593105 — —
F. dolichose aTSFC20191015-23 Republic o Ko ea ON427765 ON427795 —ON464731
F. dollinge iTDoll623 Uni ed S a es MW908540 MW898444 — —
F. eucalyp iTDai 18792 Aus alia MN816731 MN810022 MN848831 MN848778
F. eucalyp icolaTDai 18593A Aus alia PP732562 PP732631 — —
F. e ea JV 1606/2.2-J Uni ed S a es KX961100 KY189100 MH636402 MH079394
F. e uginosa Cui 9244 China MN816706 MN809995 MN848804 MN848791
F. o masonaTVRTOBFO3 B azil ON808603 ON795827 — —
F. o masona VRTO83 B azil — ON795830 — —
F. gil a JV 1209/65 Uni ed S a es MN816719 MN810006 MN848851 MN848768
F. gil oidesTSFC20180426-12 Republic o Ko ea ON427763 ON427793 ON479802 ON464729
F. hawaiiana JV 2208/H22-J Uni ed S a es OQ817709 OQ817855 OQ849746 —
F. indicaTMUBL1104 India — PP390498 PV638743 PV638735
F. indica HRS-15B India — PQ113747 PV638744 PV638736
F. insoli aTJV1208/5208 Russia MN816724 MN810016 MN848800 —
F. ka s enianaTDai 11403 China MN816717 MN810003 MN848795 MN848807
F. ko eanaTSFC20160726-93 Republic o Ko ea ON427762 ON427792 ON479801 ON464728
F. la ispo a JV 1109/482 Uni ed S a es MG008439 MG008468 MH636395 MN848799
F. licnoides URM 83001 B azil MH392561 MH407357 — —
F. minu issimaTJV 2208/H16-J Uni ed S a es OQ817711 OQ817857 OQ849748 —
F. mon ana 175856 Taiwan JX484015 JX484007 — —
F. mon icolaTDai 11860 China MG008406 MG008457 MH636390 —
F. ma quesiana URM83094 B azil MH392544 MH407343 — —
F. nadia a iana MUBL1105 India PQ098039 PP390499 PQ346367 PQ346369
F. nadia a iana SP2F2A India PQ098040 PQ113748 PQ346368 PQ346370
F. paloma i JV 1305/3-J Uni ed S a es MN816738 MN810028 — —
F. plume iaeTDai 18858 Aus alia MN816712 MN810010 MN848843 MN848769
152
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Species S ain Coun y ITS LSU e 1-α pb2
F. pul ini o misTCMW48600 Sou h A ica MH599102 MH599127 MT108960 —
F. punc a i o mis Dai 17443 China MH050755 MH050764 — —
F. amulicolaTDai 15723 China MH050749 MH050762 MN848824 MH079398
F. esupina a Dai 20455 China PP732567 PP732636 — —
F. e icula aTSFC20160115-16 Republic o Ko ea ON427761 ON427791 ON479800 ON464727
F. haba ba ina Dai 16226 China MN816743 MN810035 MN848838 MN848784
F. oseocine ea JV 1407/84 China MN816740 MN810030 MN848819 —
F. u i inc a JV 1008/25 Uni ed S a es KJ940029 KX058575 — —
F. sa ci es JV 0402/20K Venezuela MZ264225 MZ264218 — —
F. semia ida URM 82510 B azil MH392563 MH407362 — —
F. sc uposa VRTOV473 B azil ON795836 — —
F. senex Dai 15775 China MN816746 MN810038 — —
F. sep ise aTDai 12820 Uni ed S a es MG008405 MN810033 MH636394 —
F. se i e a Dai 15706 China MH050759 MH050769 MN848842 MN159391
F. sho eaeTDai 17818 Singapo e MN816735 MN810026 MN848816 —
F. sinicaTDai 15468 China MG008412 MG008459 MH636392 —
F. sinuosa Dai 20499 China MZ264227 MZ264220 — —
F. si umalaiensisTMUBL1106 India PQ098037 PP390500 PV638741 PV638737
F. si umalaiensis SRM21 India PQ098038 PQ113746 PV638742 PV638738
F. subch yseaTDai 16201 China MN816708 MN809997 MN848811 MN848796
F. sub e eaTDai 16327 China KX961098 KY053473 —MH079401
F. submu inaTDai 19655 China MZ264228 MZ264221 — —
F. sub opicaTDai 19957 China PP732565 PP732634 — —
F. e minalianaeTMUBL1107 India PQ098033 PP390501 PV638739 PV638733
F. e minalianae VM2B India PQ098034 PQ113745 PV638740 PV638734
F. o ulosa JV 1405/2 Czech Republic KX961106 KY189106 MH636405 MN159392
F. i icola He 2081 Uni ed S a es MN121829 MN121770 — —
F. wahlbe gii JV 1312/20-Kou Spain MN81672 MG008462 — —
F. yunnanensis Cui 8182 China MH050756 MN810029 —MN848789
Ou g oup
Coni e ipo ia wei ii CFS 504 Canada AY829341 AY829345 — —
Phellinidium ag ans CBS 202.90 Canada AY558619 AY059027 — —
nea ly iden ical ee opologies, wi h li le a ia ion in s a is ical suppo .
The e o e, only he ML ee is shown (Fig. 1).
The phylogene ic analyses e ealed six majo clades ep esen ing six
species complexes in Fuscopo ia, which we e consis en wi h ea lie epo s
(Fig. 1) (Chen e al. 2020, 2023a, b). Ou Indian specimens we e eco e ed
in he F. o ulosa complex, which ecei ed obus phylogene ic suppo in
bo h ML and BI analyses (92 ML/1.00 BI). Fuscopo ia si umalaiensis o med
a sis e clade o F. haba ba ina; F. indica o med a sis e axon o F. si um-
alaiensis, F. haba ba ina, F. licnoides, and F. callimo pha, while F. e mina-
lianae o med a dis inc lineage wi hin he F. o ulosa complex. The newly
iden i ied species a e cha ac e ized by e used- e lexed o pilea e basidio-
ca ps, smalle po es, sep a e skele al hyphae, he p esence o hymenial se-
ae, he absence o mycelial se ae, he p esence o cys idioles, and b oadly
ellipsoid o subglobose basidiospo es.
153
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Taxonomy
Fuscopo ia indica M. Kaliyape umal, S. Gunaseelan & K. Kezo, sp. no .
MycoBank No: MB 859271
Fig. 2
Diagnosis. Fuscopo ia indica is diagnosed by i s annual, applana e, dimidia e ba-
sidiome wi h smoo h o glab ous, indis inc ly zona e pilea su ace, homogenous
con ex , he p esence o cys idioles, and ellipsoidal, acyanophilic basidiospo es.
Figu e 1. Phylog am gene a ed om maximum likelihood analysis based on combined ITS, n LSU, pb2, and e 1-α se-
quence da a o species o Fuscopo ia wi h he ou g oups Phellinidium ag ans (CBS 202.90) and Coni e ipo ia wei ii
(CFS504). Boo s ap suppo alues ≥ 60% o ML and Bayesian pos e io p obabili ies ≥ 0.80 a e indica ed abo e he
nodes. Species complexes a e highligh ed, and newly gene a ed sequences a e shown in bold ed, blue, and pink; ype
specimens a e agged wi h “T.”
154
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Holo ype. India • Tamil Nadu, Salem Dis ic , Ye caud, on ha dwood, 23 Janu-
a y 2018, S. Gunaseelan, HRS-15A (holo ype MUBL1104). GenBank: PP390498
(n LSU); PV638735 ( pb2); PV638743 ( e 1-α).
Figu e 2. Fuscopo ia indica (holo ype MUBL1104). a. Basidiomes; b. Po e su ace; c. T ans e se sec ion o a basidiome;
d. Dissepimen edges and skele al hyphae; e. Cys idioles; . Hymenial se ae; g. Basidioles; h. Basidia; i. Basidiospo es.
Scale ba s: 5 cm (a, b); 2 cm (c); 5 μm (d–i).
155
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
E ymology. Re e s o he ype locali y, “India.”
Desc ip ion. Basidiomes annual, pilea e, applana e, sessile o imb ica e,
so , o ligh co ky when d y. Pilei dimidia e, con ex, p ojec ing up o 4.5 cm
long, 7 cm b oad, and 1 cm hick a he base. Pileal su ace yellowish b own
(5D8), b own (6E6) o da k b own (6F8), smoo h o glab ous, indis inc ly zona e.
Ma gin b own (6E5), acu e, ma gin s e ile, ligh b own (6D8), 1 mm in hickness.
Po e su ace ligh b own (6D8) o da k b own (6F8). Po es ound o angula ,
6–8 pe mm. Ma ginal se ae absen . Con ex ligh b own (6D8), homogenous,
up o 3 mm in hickness. Tube laye yellowish-b own (6D8) o b ownish-yellow
(5C7), ha d and co ky, wi h ubes up o 2 mm long.
Hyphal sys em. Hyphal sys em dimi ic; gene a i e hyphae simple sep a e;
issue da kening bu o he wise unchanged in KOH.
Con ex . Gene a i e hyphae, hyaline o pale yellow, hin o sligh ly hick-
walled, b anched, equen ly sep a e, 2–4 µm; skele al hyphae dominan , us -
b own, hick-walled wi h a medium o wide lumen, unb anched, a ely sep a e,
mo e o less s aigh , and egula ly a anged, 2.2–3.5 µm.
Tubes. Gene a i e hyphae, dominan a he dissepimen edges and sub-
hymenium, hin-walled, equen ly b anched and sep a e, hyaline o pale
yellow, 1.8–3.4 µm, some enc us ed a dissepimen edges and in hymeni-
um; skele al hyphae dominan , hick-walled wi h a medium o wide lumen,
mo e o less s aigh , subpa allel along he ubes, yellow o golden yellow,
2–3.2 µm. Hymenial se ae subula e o en icose, acu e o acumina e a he
apex, enc us ed, mos ly o igina ing om amal hyphae, da k b own, hick-
walled, 8−39 × 5−8 µm; Cys idioles hyaline o pale yellow in wa e , usoid
o subula e, a e, ape ing a he end, 6–42 × 2.8–7 µm. Basidioles cla a e
o b oadly cla a e 6.5–18 × 2.5–6.5 µm. Basidia b oadly cla a e, hyaline,
ou s e igma a, 7–17 × 2.8–6.7 µm. Basidiospo es ellipsoid, hyaline, hin-
walled, smoo h, CB−, IKI−, (3.2–)3.5–4.5(–4.8) × (2.2–)2.5–2.9(–3.2) µm,
Q=1.5, Q ange = 1.3–1.7, (n = 30/2).
Dis ibu ion. India (Tamil Nadu).
Addi ional specimen examined (pa a ype). India • Tamil Nadu, Salem
Dis ic , Ye caud (11°79'63"N, 78°21'20"E), on ha dwood, 23 Janua y 2018,
S. Gunaseelan, HRS-15B. GenBank: PQ113747 (n LSU); PV638736 ( pb2);
PV638744 ( e 1-α).
No es. The phene ic cha ac e s o F. indica and F. licnoides a e simila ,
bo h ha ing a concen ically zona e, glab ous pileal su ace, homogeneous
con ex , and acu e ma gin. Howe e , F. indica di e s in ha ing imb ica e,
dimidia e, con ex pilei and smalle basidiospo es (F. indica 3.2–4.8 × 2.2–
3.2 μm s. F. licnoides 4–5 × 2.5–3.5 μm) (Oli ei a and Gibe oni 2023). Fus-
copo ia indica di e s om F. callimo pha in ha ing an imb ica e basidiome,
con ex pilei, smalle po es, and la ge hymenial se ae (Dai 2010). Fusco-
po ia indica, F. senex, and F. o ulosa sha e simila ea u es such as appla-
na e, dimidia e basidiomes wi h a dimi ic hyphal sys em, he p esence o
cys idioles, and cyanophilic basidiospo es, bu F. indica di e s in ha ing a
glab ous, azona e pilea su ace and la ge basidiospo es. Fuscopo ia indi-
ca also di e s om F. haba ba ina by he absence o a c us abo e he con-
ex a ma u i y and, mic oscopically, by la ge basidiospo es. In con as ,
F. haba ba ina is iden i ied by he p esence o a c us abo e he con ex in
olde specimens and smalle spo es (3.2–4.2 × 2–2.5 μm) (Dai 2010).
162
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
mul ima ke phylogene ic analysis in conjunc ion wi h de ailed mo phological
obse a ions o delinea e he h ee no el species wi hin he F. o ulosa complex
collec ed om he Eas e n Gha s o Tamil Nadu. Fu he mo e, addi ional sam-
pling om he unde s udied egions o Tamil Nadu, pa icula ly he Wes e n
Gha s, is likely o enhance ou unde s anding o he axonomic complexi y and
species di e si y o Fuscopo ia.
Acknowledgemen s
Mala izhi Kaliyape umal and Sugan ha Gunaseelan hank EMR-SERB, DST
(EMR/2016/003078), and he Go e nmen o India o inancial assis ance. MK
and GS a e g a e ul o he P incipal Chie Conse a o o Fo es s, Tamil Nadu
Fo es Depa men , o g an ing pe mission (E2/20458/2017) and o hei as-
sis ance and suppo du ing ield isi s in he Eas e n Gha s o Tamil Nadu.
MK and Elango an A umugam acknowledge he Tamil Nadu S a e Council o
Highe Educa ion (RGP/2019-20/MU/HECP-0040) o inancial aid. Saman ha
C. Ka una a han hanks he Na ional Na u al Science Founda ion o China (No.
32260004), he High-Le el Talen Rec ui men Plan o Yunnan P o ince (High-
End Fo eign Expe s P og am), and he Key Labo a o y o he Yunnan P o incial
Depa men o Educa ion o he Deep-Time E olu ion on Biodi e si y om he
O igin o he Pea l Ri e o hei suppo . The au ho s ex end hei app ecia ion
o he Ongoing Resea ch Funding P og am (ORF-C -2025-6), King Saud Uni e -
si y, Riyadh, Saudi A abia. NS and JK hank Chiang Mai Uni e si y o hei sup-
po . P o . S e en L. S ephenson, Depa men o Biological Sciences, Uni e si y
o A kansas, USA, is hanked o English edi ing.
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.
Funding
This esea ch was unded by EMR-SERB, DST (EMR/2016/003078), Go e nmen o In-
dia; he Tamil Nadu S a e Council o Highe Educa ion (RGP/2019-20/MU/HECP-0040);
and he Ongoing Resea ch Funding P og am (ORF-C -2025-6), King Saud Uni e si y, Ri-
yadh, Saudi A abia.
Au ho con ibu ions
Concep ualiza ion: MK, SG, KK, EA; Da a cu a ion: MK, SG, KK, EA; Fo mal analysis: MK;
Funding acquisi ion: MK, AF, TA, AE, EC, SK; In es iga ion: MK, SG, KK, EA; Me hodology:
MK, KK, SG, EA; P ojec adminis a ion: MK; Resou ces: MK, SG, KK, EA; So wa e: MK,
KK; Supe ision: MK; Valida ion: MK, SG; Visualiza ion: MK; W i ing—o iginal d a : MK,
SG, KK, EA; W i ing— e iew and edi ing: MK, KK, SK.
163
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Au ho ORCIDs
Sugan ha Gunaseelan h ps://o cid.o g/0000-0001-7089-2292
Kezhocuyi Kezo h ps://o cid.o g/0000-0002-3723-0462
Elango an A umugam h ps://o cid.o g/0009-0003-9720-2978
Saman ha C. Ka una a hna h ps://o cid.o g/0000-0001-7080-0781
Abdallah M. Elgo ban h ps://o cid.o g/0000-0003-3664-7853
Ja u ong Kumla h ps://o cid.o g/0000-0002-3673-6541
Naka in Suwanna ach h ps://o cid.o g/0000-0002-2653-1913
Ekachai Chukea i o e h ps://o cid.o g/0000-0002-9968-5841
Mala izhi Kaliyape umal h ps://o cid.o g/0000-0002-1218-3778
Da a a ailabili y
All holo ype and pa a ype collec ions o he new species a e deposi ed a he Mad as
Uni e si y Bo any Labo a o y (MUBL), Cen e o Ad anced S udies in Bo any, Uni e si y
o Mad as, Chennai 600025, Tamil Nadu, India. The sequences gene a ed du ing his
s udy a e deposi ed in NCBI GenBank. The ITS, n LSU, pa ial pb2, and e 1-α alignmen
is deposi ed in FigSha e.
Re e ences
Bakshi BK (1971) Indian Polypo aceae (On T ees and Timbe ). Indian Council o Ag icul-
u al Resea ch, New Delhi.
Bal aza JM, T ie eile -Pe ei a L, Logue cio-Lei e C, Ry a den L (2009) San a Ca a ina
Island mang o es 3: a new species o Fuscopo ia. Mycologia 100: 859–863. h ps://
doi.o g/10.3852/08-082
Bal aza JM, Gibe oni TB (2010) New combina ions in Phellinus s.l. and Inono us s.l.
Myco axon 111: 205–208. h ps://doi.o g/10.5248/111.205
Chen Q, Dai YC (2019) Two new species o Fuscopo ia (Hymenochae ales, Basidiomy-
co a) om sou he n China based on mo phological cha ac e s and molecula e i-
dence. MycoKeys 61: 75–89. h ps://doi.o g/10.3897/mycokeys.61.46799
Chen Q, Yuan Y (2017) A new species o Fuscopo ia (Hymenochae ales, Basidiomyco-
a) om sou he n China. Mycosphe e: Jou nal o Fungal Biology 8(6): 1238–1245.
h ps://doi.o g/10.5943/mycosphe e/8/6/9
Chen Q, Wu F, Ji XH, Si J, Zhou LW, Tian XM, Vlasák J, Dai YC (2019) Phylogeny o he
genus Fuscopo ia and axonomic assessmen o he F. con igua g oup. Mycologia
111(3): 1–22. h ps://doi.o g/10.1080/00275514.2019.1570749
Chen Q, Du P, Vlasak J, Wu F, Dai YC (2020) Global di e si y and phylogeny o Fuscopo ia
(Hymenochae ales, Basidiomyco a). Mycosphe e: Jou nal o Fungal Biology 11(1):
1477–1513. h ps://doi.o g/10.5943/mycosphe e/11/1/10
Chen Q, Liu L, Zhang DS, Dong LL (2022) Fuscopo ia hainanensis sp. no . (Hymenochae-
ales, Basidiomyco a), a new membe o he F. con igua g oup. Phy o axa 558(3):
251–262. h ps://doi.o g/10.11646/phy o axa.558.3.1
Chen Q, Liu L, Si J, Vlasák J (2023a) Taxonomic and phylogene ic con ibu ions o
Fuscopo ia (Hymenochae ales, Basidiomyco a): Two new species om Hawaii wi h
a key o No h Ame ican species. F on ie s in Cellula and In ec ion Mic obiology 13:
1205669. h ps://doi.o g/10.3389/ cimb.2023.1205669
Chen Q, Luo HD, Cheng N, Zhang DS (2023b) Mo phological and molecula e idence
o a new species o Fuscopo ia (Hymenochae ales, Basidiomyco a) om opics.
Phy o axa 619(3): 219–231. h ps://doi.o g/10.11646/phy o axa.619.3.2
164
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Chen Q, Chen H, Luo CH, Lai XH (2024) Molecula phylogeny and mo phology o he genus
Fuscopo ia (Hymenochae ales, Basidiomyco a) and e eal h ee new species o he F.
e ea g oup. MycoKeys 111: 21–40. h ps://doi.o g/10.3897/mycokeys.111.126446
Chen Q, Chen H, Luo C, Zhang D (2025) A new species o Fuscopo ia (Hymenochae ales,
Basidiomyco a) om Sou hwes China by he mo phological and molecula phyloge-
ne ic analyses. New Zealand Jou nal o Bo any 17: 1–14. h ps://doi.o g/10.1080/0
028825X.2025.2517406
Cho Y, Kim D, Lee Y, Jeong J, Hussain S, Lim YW (2023) Valida ion o Fuscopo ia
(Hymenochae ales, Basidiomyco a) ITS sequences and i e new species based
on mul i-ma ke phylogene ic and mo phological analyses. IMA Fungus 14(1): 12.
h ps://doi.o g/10.1186/s43008-023-00117-6
C ous PW, Wing ield MJ, Ju je ic Z, Balasho S, Osieck ER, Ma in-Felix Y, Luangsa-A d
JJ, Mejía LC, Cappelli A, Pa a LA, Lucchini G (2024) Fungal plane desc ip ion
shee s: 1697–1780. Fungal Sys ema ics and E olu ion 14: 325–577. h ps://doi.
o g/10.3114/ use.2024.14.19
Dai YC (2010) Hymenochae aceae (Basidiomyco a) in China. Fungal Di e si y 45(1):
131–343. h ps://doi.o g/10.1007/s13225-010-0066-9
Edle D, Klein J, An onelli A, Sil es o D (2020) axmlGUI 2.0: A g aphical in e ace and
oolki o phylogene ic analyses using RAxML. Me hods in Ecology and E olu ion 12:
373–377. h ps://doi.o g/10.1111/2041-210X.13512
Fiasson JL, Niemelä T (1984) The Hymenochae ales: A e ision o he Eu opean po oid
axa. Ka s enia 24(1): 14–28. h ps://doi.o g/10.29203/ka.1984.224
Fische M (1996) On he species complexes wi hin Phellinus: Fomi ipo ia e isi ed. Myco-
logical Resea ch 100: 1459–1467. h ps://doi.o g/10.1016/S0953-7562(96)80078-5
G oposo C, Logue cio-Lei e C, Goes-Ne o A (2007) Fuscopo ia (Basidiomyco a, Hymeno-
chae ales) in sou he n B azil. Myco axon 101: 55–63.
Gunaseelan S, Kezo K, Ka una a hna SC, Yang E, Zhao C, Elgo ban AM, Tibp omma S,
Kaliyape umal M (2024) New species o T opicopo us (Basidiomyco a, Hymenochae-
ales, Hymenochae aceae) om India, wi h a key o A o-Asian T opicopo us species.
MycoKeys 102: 29–54. h ps://doi.o g/10.3897/mycokeys.102.117067
Hussain S, Al-Kha ousi M, Al-Muha abi DA, Al-Shabibi Z, Al-Balushi AH, Al-Yahya’Ei MN,
Al Saady N, Velazhahan R, Al-Sadi AM (2022) Phylogeny, dis ibu ion and ime di e -
gence o Fuscopo ia (Hymenochae aceae, Basidiomyco a) wi h he desc ip ion o a
new species om Dho a egion, sou he n pa o Oman. Phy o axa 570(2): 150–164.
h ps://doi.o g/10.11646/PHYTOTAXA.570.2.3
Ko ne up A, Wansche JH (1978) Me huen Handbook o Colou (3 d ed.). Ey e Me huen,
London.
Kuma S, S eche G, Li M, Knyaz C, Tamu a K (2018) MEGA X: Molecula e olu iona y
gene ics analysis ac oss compu ing pla o ms. Molecula Biology and E olu ion 35:
1547–1549. h ps://doi.o g/10.1093/molbe /msy096
Kuma i A, Ha sh NSK, Singh AN (2021) Hos ange o Phellinus sensu la o in India:
An o e iew. Jou nal o Applied and Na u al Science 13(1): 157–169. h ps://doi.
o g/10.31018/jans. 13i1.2510
Lan ea R, Calco B, Ho SYW, Guindon S (2012) Pa i ionFinde : Combined selec ion o
pa i ioning schemes and subs i u ion models o phylogene ic analyses. Molecula
Biology and E olu ion 29: 1695–1701. h ps://doi.o g/10.1093/molbe /mss020
Luana G, Fabiano S, Fabio G, Paolo G (2015) Compa ing isual inspec ion o ees and
molecula analysis o in e nal wood issues o he diagnosis o wood decay ungi.
Fo es y 88(4): 465–470. h ps://doi.o g/10.1093/ o es y/cp 015
165
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Ma heny PB (2005) Imp o ing phylogene ic in e ence o mush ooms wi h RPB1 and
RPB2 nucleo ide sequences (Inocybe; Aga icales). Molecula Phylogene ics and E o-
lu ion 35(1): 1–20. h ps://doi.o g/10.1016/j.ympe .2004.11.014
Mu ill WA (1907) Polypo aceae 1. No h Am Flo a 9: 1–72.
Niemelä T, Wagne T, Fische M, Dai YCh (2001) Phellopilus gen. no . and i s a ini ies
wi hin Phellinus s. la o and Inono us s. la o (Basidiomyce es). Annales Bo anici Fen-
nici 38: 51–62. h ps://www.js o .o g/s able/23726833
Nylande JAA (2004) M Model es .2. P og am dis ibu ed by he Au ho ; E olu iona y
Biology Cen e, Uppsala Uni e si y, Uppsala, Sweden.
Oli ei a VRT, Gibe oni TB (2023) New eco ds on Fomi ipo ia and Fuscopo ia (Hymeno-
chae aceae) in a eas o he A lan ic Fo es in No heas e n B azil. Lilloa 60(1): 41–
57. h ps://doi.o g/10.30550/j.lil/2023.60.1/2023.03.20
Panconesi A, San ini A, Casini N (1994) Phellinus o ulosus on Cup essus sempe -
i ens in I aly. Eu opean Jou nal o Fo es Pa hology 24(4): 238–240. h ps://doi.
o g/10.1111/j.1439-0329.1994. b00990.x
Raymundo T, Valenzuela R, Bau is a-He nández S, Esqueda M, Ci uen es J, Pacheco L
(2013) The genus Fuscopo ia (Hymenochae ales, Basidiomyco a) in Mexico. Re is a
Mexicana de Biodi e sidad 84: 50–69. h ps://doi.o g/10.7550/ mb.31604
Rehne SA, Buckley E (2005) A Beau e ia phylogeny in e ed om nuclea ITS and EF1-a
sequences: E idence o c yp ic di e si ica ion and links o Co dyceps eleomo phs.
Mycologia 97: 84–98. h ps://doi.o g/10.1080/15572536.2006.11832842
Robe V, Vu D, Ben Hadj Amo A, an de Wiele N, B ouwe C, Jabas B, Szoke S, D idi A,
T iki M, ben Daoud S, Chouchen O, Vaas L, de Cock A, S alpe s JA, S alpe s D, dos
San os FB, Li W, Wu L, Zhang R, Ma J, Zhou M, Eu wilaichi L, Ings iswang S, Hansen
K, Schoch C, Robbe se B, I inyi L, Meye W, Hawkswo h DL, Taylo JW, C ous PW
(2013) MycoBank gea ing up o new ho izons. IMA Fungus 4(2): 371–379. h ps://
doi.o g/10.5598/ima ungus.2013.04.02.16
Ronquis F, Teslenko M, an de Ma k P, Ay es DL, Da ling A, Höhna S, La ge B, Liu
L, Sucha d MA, Huelsenbeck JP (2012) M Bayes 3.2: E icien Bayesian phylogene -
ic in e ence and model choice ac oss a la ge model space. Sys ema ic Biology 61:
539–542. h ps://doi.o g/10.1093/sysbio/sys029
Saye s EW, Ca anaugh M, F isse L, P ui KD, Schneide VA, Unde wood BA, Yankie L,
Ka sch-Miz achi I (2025) GenBank 2025 upda e. Nucleic Acids Resea ch 53(D1):
D56–D61. h ps://doi.o g/10.1093/na /gkae1114
Singh AP, Kau G, Dhing a GS (2014) Di e si y o po oid mush ooms in Punjab: amily
Hymenochae aceae. P oceedings o he 8 h In e na ional Con e ence on Mush oom
Biology and Mush oom P oduc s (ICMBMP8): 83–91.
Spi in WA, Zmi o ich V, Malyshe a VF (2006) To he sys ema ics o Phellinus s.l. and
Inono us s.l. (Muc onopo aceae, Hymenochae ales). No os i Sis ema iki Nizsih Ras-
enij 40: 153–188.
Spi in V, Vlasák J, Niemelä T (2014) Fuscopo ia insoli a (Hymenochae ales, Basidiomy-
co a), a new species om Russian Fa Eas . Ann Bo Fenn 51: 403–406. h ps://doi.
o g/10.5735/085.051.0607
Vilgalys R, Hes e M (1990) Rapid gene ic iden i ica ion and mapping o enzyma ically
ampli ied ibosomal DNA om se e al C yp ococcus species. Jou nal o Bac e iology
172(8): 4238–4246. h ps://doi.o g/10.1128/jb.172.8.4238-4246.1990
Wagne T, Fische M (2001) Na u al g oups and a e ised sys em o he Eu opean po oid
Hymenochae ales (Basidiomyco a) suppo ed by nLSU DNA sequence da a. Myco-
logical Resea ch 105(7): 773–782. h ps://doi.o g/10.1017/S0953756201004257
166
MycoKeys 125: 147–166 (2025), DOI: 10.3897/mycokeys.125.168173
Sugan ha Gunaseelan e al.: Th ee new species o Fuscopo ia om India
Wagne T, Fische M (2002) P oceedings owa ds a na u al classi ica ion o he wo ld-
wide axa Phellinus s.l. and Inono us s.l., and phylogene ic ela ionships o allied gen-
e a. Mycologia 94(6): 998–1016. h ps://doi.o g/10.1080/15572536.2003.11833156
Whi e TJ, B uns TD, Lee S, Taylo J (1990) Ampli ica ion and di ec sequencing o ungal
ibosomal RNA genes o phylogene ics. In: Innis MA, Gel and DH, Sninsky JJ, Whi e
TJ (Eds) PCR p o ocols: A guide o me hods and applica ions. Academic P ess, New
Yo k, 315–322. h ps://doi.o g/10.1016/B978-0-12-372180-8.50042-1
Wu F, Zhou LW, Vlasák J, Dai YC (2022) Global di e si y and sys ema ics o Hymeno-
chae aceae wi h po oid hymenopho e. Fungal Di e si y 113(1): 1–192. h ps://doi.
o g/10.1007/s13225-021-00496-4
Yuan HS, Lu X, Dai YC, Hyde KD, Kan YH, Kušan I, He SH, Liu NG, Sa ma VV, Zhao CL, Cui
BK, Yousa N, Sun GY, Liu SY, Wu F, Lin CG, Daya a hne MC, Gibe oni TB, Conceição
LB, Ga ibay-O ijel R, Villegas-Ríos M, Salas-Lizana R, Wei TZ, Qiu JZ, Yu ZF, Phookam-
sak R, Zeng M, Paloi S, Bao DF, Abeywick ama PD, Wei DP, Yang J, Manawasinghe IS,
Ha ishchand a D, B ahmanage RS, de Sil a NI, Tennakoon DS, Ka una a hna A, Ga -
o o Y, Pem D, Zhang SN, de Aze edo San iago ALCM, Beze a JDP, Dima B, Acha ya
K, Al a ez-Manja ez J, Bahkali AH, Bha VK, B and ud TE, Bulgako TS, Campo esi E,
Cao T, Chen YX, Chen YY, De ada ha B, Elgo ban AM, Fan LF, Du X, Gao L, Gonçal es
CM, Gusmão LFP, Huan aluek N, Jadan M, Jayawa dena RS, Khalid AN, Lange E,
Lima DX, de Lima-Júnio NC, de Li a CRS, Liu JK, Liu S, Lumyong S, Luo Z-L, Ma očec
N, Ni anjan M, Oli ei a-Filho JRC, Papp V, Pé ez-Pazos E, Phillips AJL, Qiu P-L, Ren
Y, Ruiz RFC, Semwal KC, Soop K, de Souza CAF, Souza-Mo a CM, Sun L-H, Xie M-L,
Yao Y-J, Zhao Q, Zhou L-W (2020) Fungal di e si y no es 1277–1386: Taxonomic and
phylogene ic con ibu ions o ungal axa. Fungal Di e si y 104(1): 1–266. h ps://
doi.o g/10.1007/s13225-020-00461-7
