291
Expanding he genus Neoba yopsis (Calca ispo iaceae,
Hypoc eales): desc ip ions o wo new species and wo new
combina ions based on mo phological and molecula da a
Vale ii Da mos uk1, Ja ie E ayo2, Pamela Rod iguez-Flakus1, Ma in Kukwa3, Adam Flakus1
1 W. Sza e Ins i u e o Bo any, Polish Academy o Sciences, Lubicz 46, PL-31-512 K akow, Poland
2 Na a o Villoslada 16, 3° dcha., 31003 Pamplona, Na a a, Spain
3 Depa men o Plan TaxonomyandNa u eConse a ion,Facul yo Biology,Uni e si yo Gdańsk,Wi aS wosza59,PL-80-308Gdańsk,Poland
Co esponding au ho : Vale ii Da mos uk ( .da mos uk@bo any.pl)
Copy igh : © Vale ii Da mos uk 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 Neoba yopsis (Calca ispo iaceae) comp ises lichenicolous ungi wi h
b igh -colo ed pe i hecia, long, mul isep a e, ili o m ascospo es, and mononema ous
conidiopho es o dis inc i e synnema ous-like asexual mo phs. The genus was o igi-
nally desc ibed om species g owing on Loba iella spp., bu i s close mo phological
simila i y o Neoba ya long obscu ed i s phylogene ic iden i y. In his s udy, we eas-
sess he sys ema ics o Neoba ya-like axa using a combina ion o mo phological and
mul i-locus phylogene ic analyses (ITS, LSU, e 1, pb2). Ou esul s con i m ha se -
e al lichenicolous species his o ically placed in Neoba ya in ac belong o he genus
Neoba yopsis, which is closely ela ed o Calca ispo ium. We desc ibe wo new species
(Neoba yopsis e iode ma icola and N. eloschis is) and p opose wo new combina ions
(Neoba yopsis cilia is and N. pel ige ae). This wo k expands he genus Neoba yopsis
o i e ecen ly accep ed species, some o which exhibi s ong hos speci ici y and a
p ima ily Sou h Ame ican dis ibu ion. An iden i ica ion key o species o Neoba yopsis
and Neoba ya-like ungi is p o ided. Ou esul s cla i y he sys ema ics o hese mo pho-
logically simila lichenicolous ungi and p o ide a amewo k o u u e s udies on hos
speci ici y, biogeog aphy, and e olu iona y ela ionships wi hin Hypoc eales.
Key wo ds: Ascomyco a, cloud o es , di e si y, lichenicolous ungi, mul i-locus phylog-
eny, Sou h Ame ica, axonomy
In oduc ion
Lichenicolous ungi, a specialized ecological g oup associa ed wi h lichen
hos s, ep esen a ema kably di e se assemblage wi h a wide ange o mo -
phological adap a ions. They occu on all con inen s, bu comp ehensi e
s udies ha e been limi ed o ce ain egions. In nume ous cases, he lack o
molecula sequence da a o hese ungi subs an ially limi s he accu acy o
hei gene ic assignmen and impedes con iden in e ence o hei phylogene -
ic placemen . Recen s udies ha e shown ha he lichenicolous li es yle has
e ol ed independen ly mul iple imes h oughou ungal e olu iona y his o y.
Academic edi o : Tho s en Lumbsch
Recei ed:
20 Augus 2025
Accep ed:
1 Oc obe 2025
Published:
5 No embe 2025
Ci a ion: Da mos uk V, E ayo J,
Rod iguez-Flakus P, Kukwa M, Flakus
A (2025) Expanding he genus
Neoba yopsis (Calca ispo iaceae,
Hypoc eales): desc ip ions o wo new
species and wo new combina ions
based on mo phological and
molecula da a. MycoKeys 124:
291–308. h ps://doi.o g/10.3897/
mycokeys.124.169315
MycoKeys 124: 291–308 (2025)
DOI: 10.3897/mycokeys.124.169315
292
MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
Mo eo e , his g oup also exhibi s a high deg ee o mo phological plas ici y,
making axonomy based solely on mo phology pa icula ly challenging (Suija
e al. 2015, 2024; Pino-Bodas e al. 2017; Flakus e al. 2019; Da mos uk and
Flakus 2024; Diede ich e al. 2024; E ayo e al. 2024).
The genus Neoba ya, wi h he ype species Neoba ya pa asi ica, was es ab-
lished o accommoda e ungicolous species wi h py i o m, ligh -colo ed (whi e,
g een, yellow, o o ange) pe i hecia, lacking a dis inc subiculum, and linea asci
wi h long, sep a e, ili o m ascospo es (E iksson and Hawkswo h 1986). The
gene ic concep was la e expanded o include species g owing on lichens ha
sha e simila mo phological and ana omical ea u es, al hough conce ns we e
aised abou he placemen o ungicolous and lichenicolous axa in he same
genus (E ayo 2002; Candoussau e al. 2007; E ayo and Sancho 2008).
Law ey e al. (2015) ob ained molecula da a o he gene ic ype Neoba ya
pa asi ica and o he lichenicolous species Neoba ya usneae. Thei esul s
showed ha hese wo species belong o di e en amilies wi hin Hypoc eales,
Cla icipi aceae and Hypoc eaceae, espec i ely. Based on a combina ion o mo -
phological ea u es and phylogene ic e idence, he au ho s es ablished a new li-
chenicolous genus, Lichenoba ya, wi h Lichenoba ya usneae as he ype species.
Subsequen ly, Flakus e al. (2019) desc ibed ano he mono ypic lichenico-
lous genus, Neoba yopsis, o Neoba ya-like specimens g owing on Loba iella
spp. in Boli ia. This genus was de ined by i s na owly py i o m, yellowish o
o ange ascoma a; long, ili o m ascospo es; and a synnema ous-like asexual
mo ph wi h a whi e s ipe and a hyaline, pink o yellowish-o ange conidial mass.
Recen phylogene ic analyses ha e shown ha Neoba yopsis o ms a well-sup-
po ed clade sis e o he anamo phic genus Calca ispo ium, sugges ing i s
a ini y wi h he amily Calca ispo iaceae, whe eas Neoba ya belongs o Cla ic-
ipi aceae (Flakus e al. 2019; Da mos uk e al. 2025).
Fou addi ional Neoba ya species g owing on lichen hos s (N. cilia is, N. da -
winiana, N. lichenophila, and N. pel ige ae) showed mo phological simila i y o
Neoba yopsis, bu o a long ime molecula da a o hese axa we e lacking
(E ayo 2002; Candoussau e al. 2007). In his s udy, we cla i y he phylogene ic
placemen o Neoba ya cilia is and N. pel ige ae, and we desc ibe wo addi ion-
al species ha belong o he genus Neoba yopsis.
Ma e ial and me hods
Taxon sampling and mo phological s udies
This s udy is based on eshly collec ed ma e ial o lichenicolous species,
complemen ed by he ba ium specimens deposi ed a KRAM, LPB, UGDA, and
he pe sonal he ba ium o J. E ayo (hb. E ayo). Mo phological and ana omical
cha ac e s we e examined using s anda d s e eo and compound mic oscopes
(Nikon SMZ 800 and 80i DIC; Leica S9i and S9D). Ascoma a sec ions we e
p epa ed manually using a azo blade o wi h a eezing sliding mic o ome
Mic om HM 430 (The mo Fishe Scien i ic, USA) combined wi h a BFS-MP
eezing s age and a BFS-3MP con olle . Sec ions and squash moun s we e
examined in dis illed wa e , 10% KOH (K), o lac ophenol co on blue (LPCB;
Fluka, no. 61335-100ML). Pho omic og aphs o ana omical cha ac e s we e
ob ained unde ansmi ed di e en ial in e e ence con as (DIC) mic oscopy.
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MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
All measu emen s we e made in dis illed wa e o LPCB. Measu emen s (i n
> 10) a e gi en as (min.–)x
––SD–x
–+SD(–max.), whe e x is he a i hme ic mean
and SD is he s anda d de ia ion.
DNA ex ac ion, PCR ampli ica ion, and DNA sequencing
Lichen halli wi h ascoma a o lichenicolous ungi we e s o ed a –20 °C un-
il p ocessing. Ascoma a we e emo ed om he hos hallus and ca e ully
cleaned in double-dis illed wa e on a mic oscope slide unde s e ile condi ions
o emo e hos issues and o he isible impu i ies using ul a- hin weeze s
and a azo blade. DNA was ex ac ed om 4 o 8 clean ascoma a o hymenia,
depending on each specimen, using he QIAamp DNA In es iga o Ki (Qiagen,
Ge many) ollowing he manu ac u e ’s ins uc ions. Fou loci — nuclea DNA
in e nal ansc ibed space (ITS), nuclea DNA la ge subuni (LSU), a agmen
o he DNA-di ec ed RNA polyme ase II subuni wo gene ( pb2), and a agmen
o he egion coding o p o ein syn hesis elonga ion ac o 1 alpha ( e 1) –
we e gene a ed. The p ime s used and polyme ase chain eac ion (PCR) con-
di ions a e lis ed in Table 1. Ampli ica ion was pe o med in a o al eac ion
olume o 25 μl, consis ing o 3 μl (5 μl o p o ein-coding egions) o genomic
DNA empla e, 1 μl (3 μl o p o ein-coding egions) o each o wa d and e e se
p ime , 13 μl o D eamTaq PCR Mas e Mix (2×) (The mo Fishe Scien i ic, USA),
and 7 μl (1 μl o p o ein-coding egions) o double-dis illed, deionized wa e .
PCR p oduc s we e sequenced in bo h di ec ions by Mac ogen (Ams e dam,
he Ne he lands). The newly gene a ed sequences we e ca e ully checked, as-
sembled, and edi ed manually using Geneious P o 8.0 (Bioma e s L d.) and
we e deposi ed in GenBank. De ailed in o ma ion on he sequences used in his
s udy is p o ided in Table 2.
Phylogene ic analyses
All gene a ed sequences we e i s subjec ed o BLAST nucleo ide sea ches
(Al schul e al. 1990) o disca d po en ial con amina ion by un ela ed ungi.
Alignmen s o each egion we e gene a ed using MAFFT (Ka oh and S andley
2013), as implemen ed on he GUIDANCE2 web se e (Penn e al. 2010).
The single-locus phylogenies o all loci we e gene a ed ( esul s no shown)
o de ec incong uences in opology. Pa i ionFinde 2 (Lan ea e al. 2017)
was used o selec he bes pa i ion scheme o ou da ase and subs i u ion
Table 1. Loci, p ime s, and polyme ase chain eac ion (PCR) condi ions used in his s udy.
Loci PCR p ime s Sequence (5′–3′) PCR Cycles Re e ences
ITS ITS1F CTT GGT CAT TTA GAG GAA GTA A 94 °C: 5 min (94 °C: 30 s, 54 °C: 30 s, 72 °C: 30 s) × 4 cycles;
(94 °C: 30 s, 48 °C: 30s, 72 °C: 1 min) × 32 cycles. A inal
ex ension o 72 °C: 10 min
Whi e e al. (1990);
Ga des and B uns (1993)
ITS4 TCC TCC GCT TAT TGA TAT GC
LSU LROR GTA CCC GCT GAA CTT AAG C 94 °C: 5 min (94 °C: 30 s, 54 °C: 30 s, 72 °C: 30 s) × 4 cycles;
(94 °C: 30 s, 48 °C: 30s, 72 °C: 1 min) × 32 cycles. A inal
ex ension o 72 °C: 10 min
Rehne and Samuels (1994);
Vilgalys and Hes e (1990)
LR5 TCC TGA GGG AAA CTT CG
e 1 EF1-983F GCY CCY GGH CAY CGT GAY TTY AT (95 °C: 30 s, 55 °C: 50 s, 72 °C: 1 min) × 35 cycles. A inal
ex ension o 72 °C: 10 min
Rehne and Buckley (2005)
EF1-2228R AT GAC ACC RAC RGC RAC RGT YTG
pb2 RPB2-5F GAY GAY MGW GAT CAY TTY YGG 94 °C: 90 s (94 °C: 30 s, 55 °C: 90 s, 68 °C: 2 min) × 40
cycles. A inal ex ension o 68 °C: 5 min
Liu e al. (1999)
RPB2-7CR CCC ATR GCT TGY TTR CCC AT
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MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
models o each pa i ion. A single subs i u ion model was selec ed o each
egion (K80+G o ITS1 and ITS2, TIM+I+G o 5.8S, LSU, he i s codon posi ion
o e 1 and pb2, F81+I o he second codon posi ion o e 1 and pb2, TVM+G
o he hi d codon posi ion o e 1, and HKY o he hi d codon posi ion o
pb2) unde a g eedy sea ch algo i hm and he Akaike in o ma ion c i e ion
(AIC) (Lan ea e al. 2012). Fo e 1 and pb2, each codon posi ion was analyzed
as a dis inc pa i ion: he i s , second, and hi d codon posi ions. The coding
domain sequence (CDS) o he p o ein-coding egions was de ec ed using he
Augus us web ool (S anke e al. 2008). The da ase included 21 specimens
Table 2. Lis o specimens, wi h hos /subs a e, s ain/ ouche , coun y o o igin, and GenBank accession numbe s used
in phylogene ic analyses.
Species S ain / Vouche Hos / subs a e Coun y GenBank accession numbe s
ITS LSU e 1 pb2
Albomo chellophila mo chellae KUNCC 21-10005 T Mo chella sp. China OP580900 OP580862 OP585424 –
Albomo chellophila mo chellae KUNCC 21-10100 Mo chella sp. China OR420019 OR420021 OR420838 –
Calca ispo ium a buscula CBS 111.57 Russula ellea Uni ed
Kingdom
MH857665 MH869205 – –
Calca ispo ium a buscula CBS 518.66 Bole us he
Ne he lands
MH858872 MH870517 – –
Calca ispo ium a buscula CBS 900.68 T decaying aga ic Ge many MH859249 KX442598 KX442596 KX442597
Calca ispo ium co dycipi icola CGMCC 3.17904 ui ing body o
Co dyceps mili a is
China KT945001 KX442604 KX442605 KX442607
Calca ispo ium co dycipi icola CGMCC 3.17905 T ui ing body o
Co dyceps mili a is
China KT944999 KX442599 –KX442594
Calca ispo ium co dycipi icola CGMCC 3.17906 ui ing body o
Co dyceps mili a is
China – KX442592 KX442591 KX442590
Calca ispo ium guizhouense DY05042 Co dyceps sp. China PP809658 PP809662 PP823899 –
Calca ispo ium guizhouense DY05041 T Co dyceps sp. China PP124948 PP133530 PP146564 –
Calca ispo ium xyla iicola HMAS 276836 T Xyla ia sp. I aly KX442603 KX442601 KX442595 KX442600
Calca ispo ium yuanyangense YFCC 22099256 T Ophioco dyceps
nu ans
China OQ954173 OQ954171 OQ981389 OQ981390
Neoba yopsis andensis E ayo 20-11 (LPB) Loba iella pallida Boli ia MT153958 MT153987 – –
Neoba yopsis andensis Flakus 25967.1 (KRAM
L-71220.1) T
Loba iella pallida Boli ia MT153956 MT153985 PP583634 –
Neoba yopsis andensis Flakus 25967.2 (KRAM
L-71220.2)
Loba iella pallida Boli ia MT153957 MT153986 – –
Neoba yopsis cilia is Kukwa 15174b (LPB) Leucode mia
leucomelos
Boli ia PX418470 PX418477 PX438769 PX438774
Neoba yopsis cilia is E ayo 34451 (LPB) Leucode mia bo yi Boli ia PX418469 PX418476 PX438768 –
Neoba yopsis e iode ma icola Kukwa s.n. (KRAM L-75209) T E iode ma sp. Boli ia PX418471 PX418478 – –
Neoba yopsis e iode ma icola E ayo 33243 (LPB) E iode ma sp. Boli ia PX418472 PX418479 PX438770 –
Neoba yopsis pel ige ae S öckli s.n. (KRAM L-75209) Pel ige a sp. Swi ze land PX418473 PX418480 PX438771 PX438775
Neoba yopsis pel ige ae Suija s.n. (TUF095161) Pel ige a sp. Finland PX418483 – – –
Neoba yopsis pel ige ae Da mos uk 1695 (KRAM
L-75217)
Pel ige a sp. No way PX418474 PX418481 PX438772 PX438776
Neoba yopsis eloschis is Kukwa 16418 (UGDA L) T Teloschis es
la icans
Boli ia PX418475 PX418482 PX438773 PX438777
Ve icimonospo ium
di ac um
CBS 310.72 T Cocos nuci e a,
decaying lea
Papua New
Guinea
MH860483 MH872200 – –
Ve icimonospo ium ellip icum CBS 100388 T Palmae, decaying
pe iole
Pe u MH862700 MH874307 – –
Sequences ob ained in his s udy a e shown in bold. Abb e ia ions: T: ype specimen o ex- ype s ain; –: indica es una ailable da a o sequence.
295
MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
belonging o 12 species o he gene a Calca ispo ium, Neoba yopsis, and
Ve icimonospo ium (Calca ispo iaceae), as well as wo specimens o
Albomo chellophila mo chellae, which we e selec ed as he ou g oup.
Maximum likelihood (ML) analyses we e ca ied ou using a heu is ic sea ch
as implemen ed in IQ-TREE 2.1.2 on CIPRES Science Ga eway (Ronquis e
al. 2012), and 1000 ul a as boo s ap eplica es we e selec ed o es ima e
b anch suppo (Minh e al. 2020). A Bayesian in e ence (BI) phylogene ic ee
was gene a ed in M Bayes 3.2.6 on CIPRES Science Ga eway (Ronquis e al.
2012) using he pa i ions and subs i u ion models ob ained by Pa i ionFinde
2. The pos e io p obabili ies we e calcula ed by sampling ees using he Ma -
ko chain Mon e Ca lo (MCMC) app oach. Two independen pa allel uns we e
andomly s a ed, each wi h ou inc emen ally hea ed chains ( he empe a u e
pa ame e o MCMC chains was 0.15). The MCMC was allowed o un o 100
million gene a ions, sampling e e y 1000 h ee and disca ding he i s 25%
o he sampled ees as a bu n-in ac o . The analysis was s opped when he
s anda d de ia ion o spli equencies d opped below 0.01. The esul ing ML
and BI phylogene ic ees we e isualized in FigT ee 1.4.4 (h p:// ee.bio.ed.ac.
uk/so wa e/ ig ee/) and Inkscape 0.92 (h ps://inkscape.o g/). The inal ee
opology was based on he 50% majo i y- ule consensus ee om he BI anal-
ysis, wi h Bayesian pos e io p obabili ies (BPP) and maximum likelihood boo -
s ap (MLB) alues indica ed abo e b anches. B anches wi h Bayesian pos e i-
o p obabili y alues >0.97 and maximum likelihood boo s ap alues >70 we e
conside ed suppo ed.
Resul s
Phylogene ic analyses
In his s udy, 25 new sequences om six specimens o lichenicolous ungi we e
gene a ed. The mul i-gene da ase o he amily Calca ispo iaceae included
3221 cha ac e s (568 o ITS, 909 o LSU, 910 o e 1, and 834 o pb2), o which
893 we e pa simony-in o ma i e si es, 119 we e single on si es, and 1975 we e
cons an si es. The phylogene ic ee showed a simila opology om ML and
BI analyses; he e o e, he BI ee was selec ed o ep esen and discuss he
phylogene ic ela ionships among axa (Fig. 1).
In ou phylogene ic analyses, wo species o he genus Ve icimonospo i-
um clus e ed in a well-suppo ed clade (1/100) and showed a sis e ela ion-
ship o he emaining gene a o Calca ispo iaceae. The i e species o he
anamo phic genus Calca ispo ium included in he analyses clus e ed oge he
in a mode a ely suppo ed clade (0.98/75). The species Neoba ya cilia is and
N. pel ige ae, along wi h wo undesc ibed lichenicolous species (g owing on
E iode ma sp. and Teloschis es la icans) and Neoba yopsis andensis, o med
a highly suppo ed clade (1/99), which showed a sis e ela ionship o he ge-
nus Calca ispo ium. This clade comp ises ou highly suppo ed subclades,
co esponding o specimens om he same lichen hos s, as well as a single on
specimen on Teloschis es la icans. Howe e , he in e nal ela ionships wi hin
he Neoba yopsis clade emain un esol ed. Two specimens o Neoba yopsis
e iode ma icola clus e ed in a well-suppo ed clade (1/100) and showed an un-
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MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
Figu e 1. Phylogene ic ela ionships wi hin he amily Calca ispo iaceae in e ed om a Bayesian In e ence analysis
(50% majo i y- ule consensus ee) o a combined ITS, LSU, e 1, and pb2 da a se . Two specimens o Albomo chelloph-
ila mo chellae we e used as he ou g oup. The numbe s abo e b anches indica e Bayesian pos e io p obabili ies (BPP)
> 0.97 and maximum likelihood boo s ap (MLB) suppo alues > 70% espec i ely (BPP/ MLB). Sequences om ype
ma e ial a e indica ed by supe sc ip T.
Calca ispo ium
Neoba yopsis
ou g oup
Ve icimonospo ium
0.02
Albomo chellophila mo chellae KUNCC 21.10100
Albomo chellophila mo chellae KUNCC 21.10005T
Ve icimonospo ium di ac um CBS 310.72T
Ve icimonospo ium ellip icum CBS 100388T
Neoba yopsis cilia is Kukwa 15174b
Neoba yopsis cilia is E ayo 34451
Neoba yopsis pel ige ae S öckli s.n.
Neoba yopsis pel ige ae Da mos uk 1695
Neoba yopsis pel ige ae Suija s.n.
Neoba yopsis e iode ma icola E ayo 32243
Neoba yopsis e iode ma icola Kukwa s.n.T
Neoba yopsis eloschis is Kukwa 16418T
Neoba yopsis andensis E ayo 20-11
Neoba yopsis andensis Flakus 25967.2
Neoba yopsis andensis Flakus 25967.1T
Calca ispo ium a buscula CBS 900.68T
Calca ispo ium a buscula CBS 518.66
Calca ispo ium yuanyangense YFCC 22099256T
Calca ispo ium guizhouense DY05041T
Calca ispo ium guizhouense DY05042
Calca ispo ium xyla iicola HMAS 276836T
Calca ispo ium co dycipi icola CGMCC 3.17905T
Calca ispo ium co dycipi icola CGMCC 3.17906
Calca ispo ium co dycipi icola CGMCC 3.17904
1/100
1/99
1/100
1/100
1/100
0.99/87
-/72
-/61
0.98/75
0.98/97
1/99
1/100
0.99/97
1/100
1/100
1/100
1/99
0.97/83
1/100
-/54
suppo ed sis e ela ionship o he clade comp ising N. andensis and N. elo-
schis is. The ype species o Neoba ya, N. pa asi ica, belongs o Cla icipi aceae
and is phylogene ically un ela ed o he genus Neoba yopsis.
Taxonomy
Neoba yopsis Flakus, E ayo, Kukwa & Rod . Flakus
Neoba yopsis Flakus, E ayo, Kukwa & Rod . Flakus, in Flakus, E ayo, Miadlikow-
ska, Lu zoni, Kukwa, Ma u a & Rod iguez-Flakus, Plan and Fungal Sys ema -
ics 64(2): 307 (2019)
Type species. Neoba yopsis andensis Flakus, E ayo, Kukwa & Rod . Flakus.
297
MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
Neoba yopsis cilia is (E ayo) Da mos uk, E ayo, Kukwa & Flakus, comb. no .
MycoBank No: 859618
Neoba ya cilia is E ayo, Biblio h. Lichenol. 84: 74 (2002). Basionym.
Typus. Colombia • Na iño, Pas o, co egimien o El Encano, e eda S a. Isabel,
30-S lago La Cocha (Guamués), 2700 m, en cilios de Leucode mia lu escens, July
1998, J. E ayo 17386, J. Muñoz & B. Ramí ez (holo ype COL!, iso ype hb. E ayo!).
Fo de ailed desc ip ion see (E ayo 2002).
No es. This species was desc ibed om a ew locali ies in Colombia g owing
on He e ode mia s. la . (including Leucode mia spp.) (E ayo 2002), wi h subse-
quen eco ds om Boli ia and Ecuado (E ayo 2017; his s udy). Neoba yopsis
cilia is ypically g ows on he cilia o he hos , hough in some specimens i was
ound on he lowe su ace o he halli.
Specimens examined. boli ia • Dep . Chuquisaca, P o . Belisa io Boe-
o, close o Padilla be ween Nue o Mundo and San a Rosa, 18°57'12"S,
64°16'37"W, 1790 m, ansi ion be ween Boli iano-Tucumano o es s and d y
in e -Andean ege a ion, on cilia o Leucode mia leucomelos, 16 Jul. 2015, J.
E ayo 29440 (hb. E ayo, LPB) • P o . He nando Siles, 15 km wes o Mon e
Agudo, 19°48'57"S, 64°06'00"W, 1815 m, dis u bed Tucumano-Boli iano o es ,
on cilia o Leucode mia leucomelos, 20 Jul. 2015, J. E ayo 30322 (hb. E ayo,
LPB) • Dep . Cochabamba, P o . Ca asco, nea Río Ba ea Mayu close o Mon e
Punku, lowe mon ane Yungas cloud o es , 17°32'27"S, 65°16'14"W, 2553 m,
on co icolous Leucode mia leucomelos, 28 No . 2014, M. Kukwa 15158b (LPB)
• ibid., 17°31'33"S, 65°16'21"W, 2430 m, lowe mon ane Yungas cloud o es , on
hallus o He e ode mia sp. on unk, 28 No . 2014, J. E ayo 34196 (hb. E ayo)
• ibid., nea Río Lopez Mendoza, 17°30'25"S, 65°16'51"W, 2248 m, lowe mon-
ane Yungas cloud o es , on hallus o He e ode mia japonica on unk, 27 No .
2014, J. E ayo 33933 (LPB, hb. E ayo) • ibid., Way a Mayu close o Mon e Punku,
17°33'30"S, 65°16'08"W, 2750 m, on co icolous Leucode mia leucomelos, 28
No . 2014, M. Kukwa 15174b (LPB) • alle de Zongo, bosque nublado, nea me -
al b idge, 16°07'41"S, 68°05'55"W, 2450 m, on cilia o He e ode mia sp., 29 May
2011, J. E ayo 26735 (LPB) • P o . Ti aque, Pa que Nacional Ca asco, Camino
de las Nubes-Co any Al o oad, 17°17'28"S, 65°44'05"W, 4146 m, open high An-
dean ege a ion, on cilia o co icolous Leucode mia leucomelos, 2 Dec. 2014,
J. E ayo 29629 (LPB) • Dep . La Paz, P o . Bau is a-Saa ed a, Á ea Na u al de
Manejo In eg ado Nacional Apolobamba, nea la Cu a, W o ien ed alley close
o Cha azani, 15°06'30"S, 69°01'50"W, 3550 m, open a ea wi h sch ubs and Pol-
ylepis nea he i e , on cilia o Leucode mia bo yi on wigs, 14 No . 2014, J.
E ayo 35009 (hb. E ayo, LPB) • P o . F anz Tamayo, Á ea Na u al de Manejo
In eg ado Nacional Apolobamba, below Pelechuco, 14°49'08"S, 69°03'50"W,
3560 m, open a ea wi h sh ubs and Polylepis ees, on cilia o Leucode mia
bo yi on ee, 20 No . 2014, J. E ayo 34451 (hb. E ayo, LPB) • Pa que Nacion-
al y Á ea Na u al de Manejo In eg ado Madidi, below Kea a Bajo, 14°41'47"S,
69°04'10"W, 3160 m, open a ea wi h sh ubs and sca e ed ees, on cilia o
Leucode mia ulga is on unk, 18 No . 2014, J. E ayo 34418 (hb. E ayo, LPB)
• Dep . San a C uz, P o . Caballe o, El Camino de las O quídeas, 17°50'27"S,
64°41'59"W, 2510 m, Yungas cloud o es , on cilia o He e ode mia podoca pa,
17 Aug. 2012, J. E ayo 28802 (hb. E ayo, LPB).
298
MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
Neoba yopsis e iode ma icola Da mos uk, E ayo, Kukwa & Flakus, sp. no .
MycoBank No: 859616
Fig. 2
Typus. boli ia • Dep . Cochabamba, P o . Chapa e, Pa que Nacional Ca asco,
Silla oad close o Villa Tuna i, 17°06'58"S, 65°41'19"W, 1020 m, Sub-Andean
Amazon o es close o plan a ion, on apo hecia o E iode ma sp., 3 Dec. 2014,
M. Kukwa s.n. & A. Flakus (holo ype KRAM L-75209, iso ype LPB)
E ymology. Named a e he hos lichen genus, E iode ma.
Desc ip ion. Ascoma a pe i hecioid, in small g oups, supe icial, wi h
loose whi e a achnoid subiculum, wi hou s oma a, py i o m o elonga e
py i o m, collapsing by la e al pinching when d y, (250–)310–370(–410) μm
high, (160–)210–250(–300) μm wide (n = 15), smoo h, pale o ange o o -
ange. Pe i hecial wall 25–40 µm hick, no changing colo in K, composed o
wo egions: ex e nal, pale yellow egion wi h hick-walled, isodiame ic cells,
2–4 μm diam., and inne egion wi h hyaline, hin-walled, la ened cells, 5–8
× 2.5–3.5 μm. Pe iphyses 0–1 sep a e, c. 7–10 × 1.5–3 µm. Asci cylind ical o
na owly cylind ical, 8-spo ed, (120–)135–160(–185) × (4.0–)4.2–4.8(–5.3)
Figu e 2. Neoba yopsis e iode ma icola (holo ype, excep B. and F. om E ayo 33243). A, B. Ascoma a and whi e hypho-
myce ous colony on he hos hallus; C. Squashed ascoma; D. Sec ion o he ascoma; E. Asci; F. Conidia; G. Ascus apex
(in LPCB); H. Ascus wi h ascospo es. Scale ba s: 250 µm (A, B), 50 µm (C, D), 25 µm (E), 10 µm (F, H), 5 µm (G).
299
MycoKeys 124: 291–308 (2025), DOI: 10.3897/mycokeys.124.169315
Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
µm (n = 12), apex hickened wi h a conspicuous cap c. 3.5–4 µm high. Asco-
spo es h ead-like, mul isep a e (sep a ha dly isible on young ascospo es),
no cons ic ed a he sep um, hyaline, wi h ounded apical pa s, smoo h-
walled, wis ed in he ascus, (80–)95–110(–120) × (1.8–)2.0–2.2(–2.4) µm
(n = 20). Asexual s age hyphomyce ous, p esen nea he lowe pa o pe i h-
ecia, colonies e use. Conidiopho es sho , unb anched, 0–2-sep a e, hya-
line. Conidiogenous cells e minal, hyaline, hin-walled, smoo h, cylind ical o
sligh ly ape ing, phialidic, 25–40 × 1.5–2 μm. Conidia ellipsoidal o cylind i-
cal, some imes sligh ly cons ic ed in he middle, hyaline, 0-sep a e, smoo h,
sligh ly unca ed, (7.4–)8.0–9.4(–9.6) × (2.6–)2.8–3.8(–4.2) µm (n = 25).
Hos , dis ibu ion, and ecology. Neoba yopsis e iode ma icola is cu en -
ly known om a ew locali ies in he mon ane Yungas cloud o es in Boli ia
(3200–3500 m) and he Sub-Andean Amazon o es , whe e i g ows on apo he-
cia o a ely on hallus o E iode ma spp.
No es. Neoba yopsis e iode ma icola is mo phologically simila o N. pel ig-
e ae, which g ows on Pel ige a spp. (see no es below). Neoba yopsis pel ige -
ae di e s om he new species in possessing subglobose o sligh ly cla a e
e minal cells a he pe i hecial apex, a ea u e absen in N. e iode ma icola.
Addi ionally, N. pel ige ae has sho e ascospo es, 35–75 µm ( s. (80–)95–
110(–120) µm in N. e iode ma icola), and di e en lichen hos s (Candoussau
e al. 2007). The anamo phic s age o Neoba yopsis e iode ma icola also has
smalle conidia, (7.4–)8.0–9.4(–9.6) × (2.6–)2.8–3.8(–4.2) µm, compa ed o
hose epo ed om cul u es o N. pel ige ae (11–14 × 3–7 µm).
Ou phylogene ic esul s indica e ha he new species is sis e o Neo-
ba yopsis andensis and N. eloschis is, al hough his ela ionship lacks signi -
ican s a is ical suppo (Fig. 1). Neoba yopsis andensis can be dis inguished
by i s la ge ascoma a, 500–700 µm high ( s. (250–)310–370(–410) μm high
in N. e iode ma icola), longe asci and ascospo es, smalle conidia, 4–7 ×
2–2.5 μm ( s (7.4–)8.0–9.4(–9.6) × (2.6–)2.8–3.8(–4.2) µm in N. e iode ma i-
cola), as well as di e en lichen hos s (Flakus e al. 2019).
Specimens examined. boli ia • Dep . Cochabamba, P o . Ti aque, Pa que
Nacional Ca asco, Camino de los Nubes, An enas Silla -Villa Tuna i old oad,
17°12'32"S, 65°41'52"W, 3520 m, uppe mon ane Youngas cloud o es , on hal-
lus and especially apo hecia o E iode ma sp. on wigs, 30 No . 2014, J. E ayo
33243 (hb. E ayo, LPB) • Dep . La Paz, P o . No Yungas, Pa que Nacional y
Á ea Na u al de Manejo In eg ado Co apa a o Undua i by Sillu Tinca a p e-Co-
lumbian ou e, ansi ion Pá amo Yungeño – Yungas mon ane cloud o es ,
16°17'22"S, 67°53'29"W, 3518 m, on E iode ma leylandii, 25 May 2011, J. E ayo
28016 (LPB) • ibid., bosque nublado yungas, sende o que pa e de la Es ación
de se icio, 16°17'09"S, 67°51'00"W, 3220–3250 m, on E iode ma leylandii on
wigs, 24 May 2011, J. E ayo 27692 (hb. E ayo).
Neoba yopsis pel ige ae (Lowen, Boque as & Gómez-Bolea) Da mos uk,
comb. no .
MycoBank No: 859619
Neoba ya pel ige ae Lowen, Boque as & Gómez-Bolea, Sydowia 59(2): 206
(2007). Basionym.
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Vale ii Da mos uk e al.: Expanding he genus Neoba yopsis
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