205
An o e iew o he genus Lobo hallia (lichenized Ascomyco a,
Megaspo aceae) in China
Lun Wang1*, Yi-Shan Feng1*, Li-Song Wang2,3, Xin-Yu Wang2,3 , Yan-Yun Zhang1
1 Key Labo a o y o Biodi e si y Conse a ion and Ecological Secu i y in he Yang ze Ri e Basin o Anhui P o ince, College o Li e Sciences, Anhui No mal
Uni e si y, 241000 Wuhu, China
2 S a e Key Labo a o y o Phy ochemis y and Na u al Medicines, Kunming Ins i u e o Bo any, Chinese Academy o Sciences, 650201 Kunming, China
3 Yunnan Key Labo a o y o Fungal Di e si y and G een De elopmen , Kunming Ins i u e o Bo any, Chinese Academy o Sciences, 650201 Kunming, China
Co esponding au ho s: Xin-Yu Wang ([email p o ec ed]); Yan-Yun Zhang ([email p o ec ed])
Copy igh : © Lun Wang 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
Lobo hallia is a species- ich genus o c us ose lichens ha is mainly dis ibu ed in he
No he n Hemisphe e. In ecen yea s, signi ican ad ancemen s ha e been made in
he axonomy and phylogeny o his genus, leading o he desc ip ion o nume ous new
species. Howe e , Lobo hallia in China has ne e been sys ema ically e ised. In his
s udy, app oxima ely 500 specimens o Lobo hallia collec ed om di e en p o inces
o China we e examined. Based on mo phological and mul i-gene phylogene ic analy-
ses, an o e iew o his genus in China was p o ided. Eigh species, L. benzilanensis,
L. complana a, L. polypycnidia a, L. pseudop uinosa, L. pulch a, L. ub a, L. s ipi a a and
L. wangii, we e desc ibed as new o science. The species, Lobo hallia b achyloba, was
epo ed in China o he i s ime. In addi ion, we gene a ed DNA sequences o he spe-
cies Lobo hallia hedinii om opo ype specimens and cla i ied i s phylogene ic posi ion.
De ailed desc ip ions, illus a ions o mo phological cha ac e s o he abo e-men ioned
species, and compa isons wi h closely ela ed axa a e p o ided. A dicho omous key o
22 species o Lobo hallia om China is p esen ed.
Key wo ds: Lichen- o ming ungi, new species, Pe usa iales, phylogeny, species delimi a ion
In oduc ion
Acco ding o he es ima ion o Hyde e al. (2024), ungi comp ise 19 phyla,
83 classes, 1,220 amilies, 10,685 gene a and app oxima ely 140,000 species.
Licheniza ion ep esen s a majo ungal li es yle, wi h ca. 19,000 known liche-
nized species (lichens) wo ldwide (Lücking e al. 2017), including abou 3,000
epo ed om China (Wei 2020). O e he pas decades, while China has wi -
nessed a signi ican inc ease in lichenological esea ch ou pu , s udies ocus-
ing on he genus Lobo hallia (Clauzade & Cl. Roux) Ha ellne emain sca ce
(Kou e al. 2013; Pauko e al. 2019; Zhang e al. 2020, 2024a).
The axonomic concep o Lobo hallia was ini ially es ablished as a sub-
genus wi hin Aspicilia A. Massal. (Clauzade and Roux 1984), wi h subse-
quen ele a ion o gene ic ank in Aspiciliaceae (ined.) by Ha ellne (1991).
Academic edi o : Tho s en Lumbsch
Recei ed:
30 Sep embe 2025
Accep ed:
6 No embe 2025
Published:
20 No embe 2025
Ci a ion: Wang L, Feng Y-S,
Wang L-S, Wang X-Y, Zhang Y-Y
(2025) An o e iew o he genus
Lobo hallia (lichenized Ascomyco a,
Megaspo aceae) in China.
MycoKeys 125: 205–244. h ps://doi.
o g/10.3897/mycokeys.125.173554
MycoKeys 125: 205–244 (2025)
DOI: 10.3897/mycokeys.125.173554
* These au ho s con ibu ed equally o his wo k.
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MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554
Lun Wang e al.: An o e iew o he genus Lobo hallia in China
Schmi e al. (2006) econs uc ed he phylogene ic amewo k o Pe u-
sa iales using nuclea la ge subuni (nuLSU) and mi ochond ial small sub-
uni (m SSU) ibosomal DNA sequences, p oposing he ans e o Aspicilia
and Lobo hallia o Megaspo aceae Lumbsch and ega ding Aspiciliaceae as
a synonym o Megaspo aceae. No din e al. (2010) econs uc ed he phy-
logeny o Megaspo aceae by in eg a ing n LSU and m SSU da a and con-
i med he monophyly o bo h Megaspo aceae and Lobo hallia. Subsequen
esea che s ha e con inually e ised Megaspo aceae, in oducing se e al
new gene a (Soh abi e al. 2013b; Haji Moni i e al. 2017; Zake i e al. 2017;
Pauko e al. 2024; Wheele e al. 2024). To da e, ele en gene a wi hin his
amily a e accep ed (Wheele e al. 2024), wi h Lobo hallia exhibi ing he
closes phylogene ic a ini y o Teu oa Soh abi & S.D. Lea .
Lobo hallia is widely dis ibu ed in a id and moun ainous egions o he
No he n Hemisphe e, hough species ichness a ies conside ably ac oss di -
e en pa s o Eu asia (Pauko e al. 2019). A ew widesp ead axa, such as
Lobo hallia alphoplaca (Wahlenb.) Ha ellne , ha e been eco ded in he Sou h-
e n Hemisphe e, o example in New Zealand (Galloway and Ledingham 2012).
As he hi d mos species- ich genus wi hin Megaspo aceae, Lobo hallia cu -
en ly encompasses 31 accep ed species: L. alphoplaca, L. b achyloba Pauko &
I.V. F olo , L. ce noho skyana (Clauzade & Vězda) A. No din e al., L. chade audiana
(Cl. Roux) A. No din e al., L. che esina (Müll. A g.) A. No din e al., L. con o e sa
Cl. Roux & A. No din, L. c assima gina a Kou & Q. Ren, L. c enula a Lun Wang &
Y.Y. Zhang, L. densip uinosa A. Ash a e al., L. de e mina a (H. Magn.) T.B. Wheel-
e , L. elobula a Zul iqa e al., L. epiadelpha Pauko & A. No din, L. gangwondoana
S.Y. Kond . e al., L. hedinii (H. Magn.) Pauko e al., L. hyd ocha is (Poel & Nimis)
Soh abi & Nimis, L. iqbalii Zul iqa e al., L. kumino ae (Sedeln.) Sedeln., L. lac eo-
la (Oxne ) Şenka d. e al., L. lobula a Lun Wang & Y.Y. Zhang, L. melanaspis (Ach.)
Ha ellne , L. pakis anica Razzaq e al., L. pel as ic oides (Hasse) T.B. Wheele ,
L. pla yca pa (J. S eine ) Pauko , L. p ae adiosa (Nyl.) Ha ellne , L. p uinosa Kou
& Q. Ren, L. pul ina a R. Zul iqa e al., L. adiosa (Ho m.) Ha ellne , L. ecedens
(Taylo ) A. No din e al., L. semis e ilis (H. Magn.) Y.Y. Zhang, L. subdi ac a (H.
Magn.) Pauko and L. zog ii Pauko & Da ydo (Ha ellne 1991; No din e al.
2010; Roux 2012; Kou e al. 2013; Nimis 2016; Roux e al. 2016; Sedelniko a and
Sedelniko 2018; Pauko e al. 2019; Kond a yuk e al. 2020; Sipman and Ap oo
2020; Zhang e al. 2020, 2024a; Ash a e al. 2022; Zul iqa e al. 2022, 2023;
Nascimbene e al. 2023; Wheele e al. 2024). The axonomic nomencla u e o
se e al species wi hin he genus has unde gone e ision. Lobo hallia a inosa
(Flö ke) A. No din, Sa ić & Tibell, o iginally p oposed by No din e al. (2010) as
a new combina ion based on he in alid basionym Aspicilia a inosa sensu Nyl.,
was subsequen ly eclassi ied by Roux e al. (2016) as he no el species Lobo h-
allia con o e sa. Lobo hallia pa asi ica (B. de Lesd.) May ho e e al. (May ho e
e al. 2005) is conside ed a synonym o L. adiosa, since i s basionym, Aspicilia
pa asi ica B. de Lesd., was ea ed as Lobo hallia adiosa chemo ype pa asi ica
by Pauko e al. (2019) based on iso ype specimen s udy. Kou e al. (2013) de-
sc ibed Lobo hallia helanensis Kou & Q. Ren om Nei Mongol, China; howe e ,
subsequen phylogene ic and mo phological analyses by Pauko e al. (2019)
demons a ed ha Lecano a subdi ac a H. Magn. and Lobo hallia helanensis
a e conspeci ic. Since he epi he “subdi ac a” has nomencla u al p io i y, Lo-
bo hallia helanensis was ecombined as L. subdi ac a (H. Magn.) Pauko .
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MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554
Lun Wang e al.: An o e iew o he genus Lobo hallia in China
In China, Lobo hallia p edominan ly occu s in he a id and moun ainous en i-
onmen s o no he n and no hwes e n egions, as documen ed in bo h his o ical
and con empo a y s udies (Magnusson 1940, 1944; Mamu e al. 2010, 2012; Kou
e al. 2013; Pauko e al. 2019; Zhang e al. 2020, 2024a). P io o his s udy, 13
species we e known om China, wi h 10 species (i.e., L. c assima gina a, L. che -
esina, L. c enula a, L. de e mina a, L. hedinii, L. lobula a, L. p uinosa, L. semis e ilis,
L. subdi ac a, and L. zog ii) ha ing hei ype locali ies wi hin he coun y. Du ing
ou in es iga ion o lichen di e si y in China, app oxima ely 500 Lobo hallia spec-
imens we e examined and o e 450 sequences (ITS, n LSU, m SSU) we e gen-
e a ed. Using an in eg a i e app oach combining mo phology, ana omy, chemis-
y, and mul i-locus phylogene ic econs uc ion, we conduc ed comp ehensi e
molecula phylogene ic analyses and species delimi a ion wi hin he genus. This
in es iga ion esul ed in he disco e y o eigh new species and one new eco d
o China, while also cla i ying he phylogene ic posi ion o Lobo hallia hedinii o
he i s ime. De ailed mo phological desc ip ions wi h mic oscopic pho og aphs
a e p o ided o hese en species, accompanied by in e speci ic compa a i e
discussions. A dicho omous key o all Chinese Lobo hallia species is p esen ed.
Ma e ials and me hods
Sample collec ion, mo phology and chemical examina ion
Th ough ieldwo k, loans, and he ba ium isi s, app oxima ely 500 Lobo hallia
specimens we e examined in his s udy. These specimens we e collec ed om
nine p o inces in China, namely Gansu, Hebei, Nei Mongol, Ningxia, Qinghai, Sich-
uan, Xizang, Xinjiang and Yunnan, co e ing he main dis ibu ion a eas o his ge-
nus in he coun y. Vouche specimens a e deposi ed in he ollowing he ba ia: Bo -
any He ba ium, College o Li e Sciences, Anhui No mal Uni e si y (AHUB), Lichen
He ba ium, Kunming Ins i u e o Bo any, Chinese Academy o Sciences (KUN-L)
and He ba ium, College o Li e Sciences, Shandong No mal Uni e si y (SDNU).
High- esolu ion images o ype specimens o some species we e p o ided by he
Swedish Museum o Na u al His o y (S) and he Koma o Bo anical Ins i u e o
RAS (LE) o ob ained om he websi e o Global Plan s (h ps://plan s.js o .o g/).
Specimens we e examined using s anda d mic oscopy echniques. Ex e nal
mo phological cha ac e s we e obse ed on ai -d ied ma e ial unde a Nikon
SMZ 745T (Mina o Ci y, Japan) s e eomic oscope. Ana omical ea u es we e
s udied using a Zeiss Axio Scope A1 ligh mic oscope (Obe kochen, Ge many).
C oss-sec ions o apo hecia and halli we e cu by hand wi h a azo blade and
obse ed a e moun ing in wa e , 10% aqueous solu ion o po assium hyd ox-
ide (K), 10% wa e solu ion o ni ic acid HNO3 (N), Lugol’s iodine solu ions (I),
o lac ophenol co on blue (LCB). Lichen subs ances we e p elimina ily exam-
ined by spo es s using he ollowing eagen s: K, C (sodium hypochlo i e solu-
ion), KC (C a e p e ea men wi h K), and P (1,4-p-phenylenediamine). Fu he
iden i ica ion was conduc ed by hin laye ch oma og aphy (TLC) using sol en
sys em C (O ange e al. 2001; Elix 2014).
In Lobo hallia, he subhymenium and hypo hecium a e ypically poo ly delim-
i ed, e en in sec ions s ained wi h LCB. To a oid con usion, we he e o e adop
he e m “subhymenial laye s” o he a ea be ween he lowe limi o he hy-
menium and he base o he apo hecium (I ano ich e al. 2025). The sepa a e
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MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554
Lun Wang e al.: An o e iew o he genus Lobo hallia in China
plec enchyma a op he hymenium is called epi hecium, which is di e en om
he pigmen ed laye known as epihymenium (Bunga z 2002). Ascospo e mea-
su emen s a e p esen ed as: (lowes eco ded–) (x
– - SD) – x
– – (x
– + SD) (–high-
es eco ded), whe e x
– is he a i hme ic mean and SD is he s anda d de ia ion,
ollowed by he numbe o measu emen s (n). The leng h- o-wid h a ion o as-
cospo es is indica ed as “l/w”.
DNA ex ac ion, PCR ampli ica ion and sequencing
Genomic DNA was ex ac ed om d ied specimens using he DNA secu e Plan
Ki (Tiangen, Beijing, China). PCR ampli ica ion was pe o med wi h he p ime s
ITS1F and ITS 4a (Ga des and B uns 1993; La ena e al. 1999) o he ITS egion;
LR0R and LR5 (Vilgalys and Hes e 1990; Rehne and Samuels 1994) o he
LSU/28S DNA egion; and m SSU1 and m SSU3R (Zolle e al. 1999) o he
mi ochond ial small subuni (m SSU) RNA egion. Polyme ase chain eac ion
(PCR) was ca ied ou in a 25 μL olume, con aining 12.5 μL 2 × Taq Mas e Mix
(Aidlab, Hong Kong), 1 μL o each p ime , 9.5 μL ddH2O and 1 μL DNA. The he -
mal cycling condi ions ollowed Zhang e al. (2024b). PCR p oduc s we e se-
quenced by Sangon Bio ech (Shanghai, China) using he ampli ica ion p ime s.
Sequence alignmen and phylogene ic analyses
The aw sequences we e ini ially e i ied using he BLAST ool on he NCBI
online se ice (h ps://blas .ncbi.nlm.nih.go /Blas .cgi) o con i m he lichen
a ini y. Fo wa d and e e se sequences we e assembled in Geneious P ime
2024.1 (h ps://www.geneious.com) and manually edi ed. We used newly gen-
e a ed n ITS, n LSU and m SSU sequences and addi ional sequences ob ained
om GenBank (Table 1) o cons uc a phylogene ic ee. The newly gene a ed
sequences ha e been deposi ed in GenBank.
Ma ixes o n ITS, n LSU, m SSU we e aligned sepa a ely using MAFFT . 7
(h ps://ma .cb c.jp/alignmen /se e /) wi h he E-INS-I s a egy (Ka oh and
S andley 2013; Ka oh e al. 2019) unde de aul pa ame e s. The alignmen s
we e checked in MEGA . 7 (Kuma e al. 2016) and mino misalignmen s we e
manually adjus ed. Subsequen ly, Maximum Likelihood (ML) single-locus ees
we e cons uc ed in IQT ee (Nguyen e al. 2015; T i inopoulos e al. 2016) and
compa ed o assess po en ial, s ongly suppo ed con lic s among gene ees.
When all s a is ically suppo ed clades (boo s ap suppo alues≥70%) we e
consis en ac oss indi idual gene ees, he da ase s we e conca ena ed. The
h ee loci we e combined ia he Conca ena e Sequence unc ion in Phylo-
Sui e 1.2.3 (Xiang e al. 2023). ModelFinde (Kalyaanamoo hy e al. 2017)
was used o es ima e he bes schemes and nucleo ide subs i u ion models
o maximum likelihood (ML) and Bayesian in e ence (BI) analyses. The bes
schemes and selec ed models a e shown in Table 2.
Bayesian ee in e ence was ca ied ou by M Bayes 3.2 (Ronquis e al.
2012). Two independen uns o ou Ma ko Chain Mon e Ca lo (MCMC) we e
pe o med each o 5 M gene a ions, sampling e e y 1000 gene a ion. To as-
sess con e gence, he a e age s anda d de ia ion o spli equencies (ASDSF)
among uns was calcula ed e e y 200 gene a ions, wi h he i s 25% o sampled
ees disca ded as bu n-in. The analyses we e e mina ed au oma ically when
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Lun Wang e al.: An o e iew o he genus Lobo hallia in China
Table 1. Vouche in o ma ion and GenBank accession numbe s o sequences used in his s udy. Newly gene a ed se-
quences a e in bold. ‘na’ indica es he e is no sequence a ailable.
Taxa Locali y Vouche n ITS n LSU m SSU
Aspicilia angelica Mon ana, USA Wheele 3535 PP965029 PP965095 PP971393
Aspicilia boykinii Mon ana, USA Wheele 7263 PP975030 PP965098 PP971376
Aspicilia cine ea Finnma k, No way Wheele 6277 MW447391 PP965054 MW424811
Aspicilia epiglyp a Väs e gö land, Sweden No din 6303 (UPS) EU057907 HM060756 HM060718
Aspicilia umosa Mon ana, USA Wheele 6802 PP965000 PP965068 PP971354
Aspicilia indissimilis Sweden No din 5943 (UPS) EU057909 HM060746 HM060708
Aspicilia oli aceob unnea A izona, USA Owe-La sson 8771 (ASU), ype PP965013 PP965084 PP971361
Aspicilia sub adians Alaska, USA Wheele 5070 PP965009 PP965080 PP971357
Aspilidea my inii Jäm land, Sweden No din 7447 (WIS-0079196) MW899516 MW907112 MW907106
Aspilidea my inii Sweden No din 6205 (UPS) na HM060754 HM060716
Aspilidea subadunans Alaska, USA Wheele 4628 MW899518 MW907114 MW907108
Aspilidea subadunans Alaska, USA Wheele 3942 MW899517 MW907113 MW907109
Aspiciliella cup eoglauca G eece Sipman & Raus 61847 (B) KY618843 KY576954 KY576930
Aspiciliella in e mu ans A menia Zake i 40503 (GLM) KY596011 KY576947 KY576923
Aspiciliella in e mu ans Czech Republic Palice 11394 (PRA) MH248855 MH255580 MH248888
Aspiciliella po osan ana Po ugal Sipman 63019 (B), ype KY618852 KY576962 KY576939
An idea b ucei Cali o nia, USA Owe-La sson 9161 (ASU), iso ype PP965011 PP965082 PP971359
An idea b ucei Cali o nia, USA Knudsen 15069 (ASU) PP975035 PP978599 PP974318
A os elia magni ica Republic o Ty a, Russia Da ydo 21959 (LE), iso ype PP832009 PP832002 PP842134
A os elia magni ica Republic o Ty a, Russia Da ydo 21961 (hb. Da ydo ), pa a ype PP832010 PP832003 PP842135
Ci cina ia a ida Mon ana, USA Wheele 5819 PP964998 PP978597 PP971346
C. caesiocine ea Uppland, Sweden Tibell 22612 (UPS) EU057897 HM060731 HM060693
C. esculen a As akhan, Russia Owe-La sson 9796 (UPS) JQ797511 JQ797493 JQ797485
C. gibbosa Uppland, Sweden No din 5878 (UPS) EU057908 HM060740 HM060702
C. ho mannii Mon ana, USA Wheele 3634 PP964982 PP965052 PP971331
C. lep osescens Sweden No din 5906 (UPS) EU057911 HM060749 HM060711
Lobo hallia alphoplaca Uk aine SK A20 (KW) KT456207 na KT456211
L. alphoplaca No way O–L–200411 MK812484 na na
L. benzilanensis China, Yunnan 18-60350 (KUN-L) PV956002 PV955950 na
L. b achyloba Russia, Republic o Al ai F olo 357 (UFU), holo ype MK347506 na MK348228
L. b achyloba China, Xinjiang 22-72815 (KUN-L) PV956003 na na
L. ce noho skyana I an, Sou h Kho asan Ta i 2311 (B) na JQ797496 JQ797481
L. che esina USA, U ah Wheele 7216 (hb. Wheele ) PP965022 na PP971366
L. complana a China, Sichuan 20-67554 (KUN-L), holo ype PV956004 PV955951 PV956066
L. complana a China, Sichuan 20-66561 (KUN-L) PV956005 na PV956067
L. con o e sa F ance, Rhône-Alpes Roux 25286 (UPS), holo ype na HM060761 HM060723
L. c assima gina a China, Nei Mongol Wang 20122565 (SDNU), holo ype JX476026 na na
L. c assima gina a China, Nei Mongol Tong 20122583 (SDNU) KC007439 na na
L. c enula a China, Xizang ZYY22-301 (KUN-L), holo ype PP663142 PV955952 PP663165
L. c enula a China, Xizang ZYY22-331 (AHUB) PP663141 PV955953 PP663164
L. densip uinosa Pakis an LAH 36790, holo ype MZ871507 na na
L. densip uinosa Pakis an LAH 36951 MZ871515 na na
L. de e mina a USA, Mon ana Wheele 3641 (hb. Wheele ) PP965020 na PP971365
L. de e mina a USA, Mon ana Wheele 6017 (hb. Wheele ) PP965021 na PP971398
L. elobula a Pakis an LAH37153, holo ype ON384441 na na
L. elobula a Pakis an LAH37154 ON428667 na na
L. epiadelpha Russia, O enbu g Oblas Pauko 1881 (UFU L-3189), holo ype MK347505 na MK348232
L. hedinii China, Gansu 18-59518 (KUN-L) PV956006 na na
L. hedinii China, Qinghai 20-68298 (KUN-L) PV956008 na PV956069
L. hedinii China, Xizang 22-71155 (KUN-L) PV956016 PV955960 PV956077
L. hedinii China, Qinghai 20-68147 (KUN-L) PV956019 PV955962 PV956080
L. hyd ocha is I aly, Sa dinia JN72085b (BOLO) OQ073922 na na
L. hyd ocha is I aly, Sa dinia SMNS-STU-F-0002807 (STU) OQ073923 na na
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Lun Wang e al.: An o e iew o he genus Lobo hallia in China
Taxa Locali y Vouche n ITS n LSU m SSU
L. iqbalii Pakis an LAH37149, holo ype ON384444 na na
L. iqbalii Pakis an LAH37150 ON384445 na na
L. lobula a China, Sichuan ZYY22-819 (KUN-L), holo ype PP663143 PV955963 PP663166
L. lobula a China, Sichuan ZYY22-829 (AHUB) PP663147 PV955964 PP663170
L. melanaspis Sweden, Jäm land No din 6622 (UPS) HQ259272 HM060726 HM060688
L. melanaspis No way Owe-La sson 8943a (UPS) JF825524 na na
L. pakis anica Pakis an LAH37137, holo ype ON392718 na na
L. pakis anica Pakis an LAH37139 ON392720 na na
L. pel as ic oides USA, Cali o nia Knudsen 14420 (UCR) PP965019 PP965087 PP971367
L. polypycnidia a China, Qinghai XY20-3034 (KUN-L), holo ype PV956020 PV955965 PV956081
L. polypycnidia a China, Qinghai XY20-808 (KUN-L) PV956021 PV955966 PV956082
L. polypycnidia a China, Qinghai 20-68413 (KUN-L) PV956022 na PV956083
L. polypycnidia a China, Qinghai 20-67303 (KUN-L) PV956023 na PV956084
L. polypycnidia a China, Qinghai 20-67134 (KUN-L) PV956024 PV955967 PV956085
L. polypycnidia a China, Qinghai 20-68672 (KUN-L) PV956025 na PV956086
L. polypycnidia a China, Qinghai 20-67160 (KUN-L) PV956026 PV955968 PV956087
L. polypycnidia a China, Xizang 20-68878 (KUN-L) PV956027 PV955969 na
L. p ae adiosa China, Xinjiang ZYY22-570 (AHUB) PP663151 PV955970 PP663173
L. p ae adiosa China, Xinjiang ZYY22-596 (AHUB) PP663150 PV955971 PP663172
L. p ae adiosa Russia, O enbu g oblas Pauko (UFU L-1264) MK347501 na MK348229
L. p uinosa China, Qinghai 22-73170 (KUN-L) PV956030 PV955972 PV956090
L. p uinosa China, Nei Mongol Wang 20123278 (SDNU), holo ype JX476028 na na
L. pseudop uinosa China, Xizang 19-64075 (KUN-L), holo ype PV956031 PV955973 na
L. pseudop uinosa China, Qinghai 20-68331 (KUN-L) PV956032 PV955974 na
L. pseudop uinosa China, Qinghai XY22-1089 (KUN-L) PV956033 PV955975 na
L. pseudop uinosa China, Qinghai 22-72438 (KUN-L) PV956034 PV955976 PV956091
L. pseudop uinosa China, Xizang ZYY22-324 (KUN-L) PV956035 PV955977 na
L. pseudop uinosa China, Qinghai 20-67831 (KUN-L) PV956036 PV955978 PV956092
L. pulch a China, Xizang 19-66027 (KUN-L), holo ype PV956037 na PV956093
L. pulch a China, Qinghai XY20-3169 (KUN-L) PV956038 na PV956094
L. pul ina a Pakis an CH-93, pa a ype OP268463 na na
L. pul ina a Pakis an PN-04 OP268468 na OQ772296
L. adiosa Sweden No din 5889 (UPS) JF703124 na na
L. adiosa Czech Republic, Sou he n Mo a ia Kocou ko a 10226/1 ON447612 ON391443 ON367885
L. adiosa Swi ze land Lumbsch 9 Aug. 2004 (F) na DQ780306 DQ780274
L. adiosa China, Xinjiang 22-72256 (KUN-L) PX061735 na na
L. ecedens Po ugal Sipman 62857 MN586980 na na
L. ecedens Sweden No din 6035 (UPS) HQ406807 na na
L. ub a China, Xinjiang 22-72301 (KUN-L) PV956039 PV955979 PV956095
L. ub a China, Xinjiang XY22-897 (KUN-L), holo ype PV956040 na PV956096
L. ub a China, Xinjiang 22-72886 (KUN-L) PV956041 na PV956097
L. semis e ilis China, Gansu ZYY22-704 (AHUB) PP663161 PV955980 PP663180
L. semis e ilis China, Gansu ZYY22-715 (AHUB) PP663159 PV955981 PP663178
L. subdi ac a Russia, Republic o Al ai F olo 178-1 (UFU) MK347503 na MK348233
L. subdi ac a China, Xinjiang ZYY22-628 (AHUB) PP663162 na PP663181
‘L. helanensis’ China, Nei Mongol D.B. Tong 20122518 (SDNU) JX476030 na na
‘L. helanensis’ China, Nei Mongol D.B. Tong 20122791 (SDNU) JX476031 na na
L. subdi ac a a . imosa China, Xinjiang ZYY22-647 (KUN-L) PP663156 PV955982 PP663175
L. subdi ac a a . imosa China, Xinjiang Wang e al. 22-72975 (KUN-L) PP663155 na PP663174
L. s ipi a a China, Xinjiang 22-72871 (KUN-L), holo ype PV956044 na PV956100
L. s ipi a a China, Xinjiang ZYY22-616 (KUN-L) PV956045 PV955983 PV956101
L. s ipi a a China, Xinjiang ZYY22-617 (KUN-L) PV956046 PV955984 PV956102
L. s ipi a a China, Xinjiang XY22-887 (KUN-L) PV956047 PV955985 PV956103
L. wangii China, Xizang 22-71215 (KUN-L), holo ype PV956048 na na
L. wangii China, Xizang ZYY22-334 (KUN-L) PV956049 PV955986 PV956104
L. wangii China, Xizang 19-65657 (KUN-L) PV956050 na PV956105
211
MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554
Lun Wang e al.: An o e iew o he genus Lobo hallia in China
Taxa Locali y Vouche n ITS n LSU m SSU
Megaspo a imiso edia a Albe a, Canada Sp ibille WP642 PP964997 PP978600 na
M. e ucosa An a c ica Zuo NJ2018074 (hb. unknown) OL331496 OL331779 na
M. e ucosa Eas Aze baijan, I an Sipman 55434 (B) na JQ797498 JQ797483
Och olechia subpallescens USA Lumbsch 19900a (MIN) na GU980985 GU980978
Och olechia upsaliensis USA Lumbsch 19916e (MIN) na GU980986 GU980979
Och olechia yasudae na AFTOL 882 na DQ986776 DQ986902
Oxne ia ia haey enii Sweden No din 5997 (UPS) na HM060755 HM060717
Oxne ia ia mashiginensis Sweden No din 5790 (UPS) EU057912 HM060732 HM060694
Oxne ia ia pe mu a a Sweden No din 6027 (UPS) EU057918 HM060747 HM060709
Oxne ia ia supe egens Sweden No din 6023 (UPS) EU057938 HM060751 HM060713
Sagedia mas uca a Finnma k, No way Wheele 6467 PP975034 PP965066 PP971347
S. simoensis T oms, No way Owe-La sson 9000 (UPS) EU057926 HM060739 HM060701
S. simoensis Finnma k, No way Wheele 6288 PP975033 PP965067 PP971348
S. zona a T oms, No way Owe-La sson 8942 (UPS) EU057946 HM060738 HM060700
Teu oa junipe icola U ah, USA Wheele 7390 PP965018 na PP971364
T. junipe icola U ah, USA Lea i 742 (BRY) JX306744 JX306758 na
T. uxo is Cas illa, Spain Rico & Piza o 3622 (H) JX306743 JX306757 na
T. uxo is Cas illa, Spain Rico & Piza o 765 (BRY) JX306745 JX306759 na
he ASDSF alues ell below 0.01, indica ing ha s a iona i y had been eached.
ML ee analyses we e pe o med using IQ-TREE (Nguyen e al. 2015; T i inopou-
los e al. 2016) unde de aul se ings, wi h b anch suppo e alua ed h ough
1000 s anda d non-pa ame ic boo s ap eplica es. T ees we e isualized and
g aphically imp o ed using In e ac i e T ee o Li e webpage (iTOL) (Le unic and
Bo k 2021) and Adobe illus a o 2020 SP.
Resul s
Phylogene ic analysis
A phylogene ic ee o Lobo hallia was econs uc ed based on a conca e-
na ed ma ix o n ITS, n LSU, and m SSU sequences. Och olechia subpalles-
cens Ve seghy, O. upsaliensis (L.) A. Massal., and O. yasudae Vain. om he
amily Och olechiaceae R.C. Ha is ex Lumbsch & I. Schmi we e selec ed as
ou g oups (Wheele e al. 2024). The da ase comp ised 127 samples, 83 o
which belonged o he genus Lobo hallia, including 20 ype specimens and
45 newly sequenced samples (Table 1). The conca ena ed alignmen con-
ained 118 n ITS (36 newly gene a ed), 78 n LSU (30 new), and 91 m SSU (26
new) sequences, wi h a o al leng h o 2342 bp pa i ioned in o i e egions
(Table 2). Phylogene ic econs uc ion was pe o med unde Maximum Like-
lihood (ML) and Bayesian In e ence (BI) me hods. Bo h me hods p oduced
cong uen ee opologies; he BI ee is p esen ed in he main ex (Fig. 1).
Table 2. The bes schemes and nucleo ide subs i u ion models selec ed by ModelFinde
o he conca ena ed 3-gene da ase .
Pa i ion Regions Posi ions Model o IQ-TREE Model o Bayes
Pa i ion 1 ITS1 1–238 TIM2e+I+G4 GTR+F+I+G4
Pa i ion 2 5.8S 239–395 TNe+I+G4 K2P+I
Pa i ion 3 ITS2 396–575 TIM2e+I+G4 GTR+F+I+G4
Pa i ion 4 n LSU 576–1669 GTR+F+I+G4 GTR+F+I+G4
Pa i ion 5 m SSU 1670–2342 GTR+F+I+G4 GTR+F+I+G4
212
MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554
Lun Wang e al.: An o e iew o he genus Lobo hallia in China
Och olechia subpallescens Lumbsch 19900a USA
Och olechia yasudae AFTOL 882
Och olechia upsaliensis Lumbsch 19916e USA
Aspilidea my inii No din 6205 Sweden
Aspilidea my inii No din 7447 Sweden
Aspilidea subadunans Wheele 3942 USA
Aspilidea subadunans Wheele 4628 USA
An idea b ucei Owe-La sson 9161 USA
An idea b ucei Knudsen 15069 USA
Teu oa junipe icola Lea i 742 USA
Teu oa junipe icola Wheele 7390 USA
Teu oa uxo is Rico & Piza o 765 Spain
Teu oa uxo is Rico & Piza o 3622 Spain
Lobo hallia adiosa 22-72256(KKUN-L) China, Xinjiang
Lobo hallia adiosa Kocou ko a 10226/1 Czech Republic
Lobo hallia adiosa Lumbsch 9 Aug. 2004 Swi ze land
Lobo hallia adiosa No din 5889 Sweden
Lobo hallia hyd ocha is JN72085b I aly
Lobo hallia hyd ocha is SMNS-STU-F-0002807 I aly
Lobo hallia lobula a ZYY22-819 (KUN-L) China, Sichuan Holo ype
Lobo hallia lobula a ZYY22-829 (AHUB) China, Sichuan
Lobo hallia ecedens Sipman 62857 Po ugal
Lobo hallia ecedens No din 6035 Sweden
Lobo hallia ce noho skyana Ta i 2311 I an
Lobo hallia pel as ic oides Knudsen 14420 USA
Lobo hallia c enula a ZYY22-301 (KUN-L) China, Xizang Holo ype
Lobo hallia c enula a ZYY22-331 (AHUB) China, Xizang
Lobo hallia de e mina a Wheele 3641 USA
Lobo hallia de e mina a Wheele 6017 USA
Lobo hallia con o e sa Roux 25286 F ance Holo ype
Lobo hallia che esina Wheele 7216 USA
Lobo hallia wangii 22-71215 (KUN-L) China, Xizang Holo ype
Lobo hallia wangii ZYY22-334 (KUN-L) China, Xizang
Lobo hallia wangii 19-65657 (KUN-L) China, Xizang
Lobo hallia pseudop uinosa 19-64075 (KUN-L) China, Xizang Holo ype
Lobo hallia pseudop uinosa ZYY22-324 (KUN-L) China, Xizang
Lobo hallia pseudop uinosa 20-67831 (KUN-L) China, Qinghai
Lobo hallia pseudop uinosa 22-72438 (KUN-L) China, Qinghai
Lobo hallia pseudop uinosa XY22-1089 (KUN-L) China, Qinghai
Lobo hallia pseudop uinosa 20-68331 (KUN-L) China, Qinghai
Lobo hallia densip uinosa LAH 36790, Pakis an Holo ype
Lobo hallia densip uinosa LAH 36951, Pakis an
Lobo hallia p uinosa 22-73170 (KUN-L) China, Qinghai
Lobo hallia p uinosa Wang 20123278 China, Inne mongolia Holo ype
Lobo hallia complana a 20-66561 (KUN-L) China, Sichuan
Lobo hallia complana a 20-67554 (KUN-L) China, Sichuan Holo ype
Lobo hallia ub a 22-72301 (KUN-L) China, Xinjiang
Lobo hallia ub a 22-72886 (KUN-L) China, Xinjiang
Lobo hallia ub a XY22-897 (KUN-L) China, Xinjiang Holo ype
Lobo hallia p ae adiosa ZYY22-596 (AHUB) China, Xinjiang
Lobo hallia p ae adiosa Pauko (UFU L-1264) Russia
Lobo hallia p ae adiosa ZYY22-570 (AHUB) China, Xinjiang
Lobo hallia semis e ilis ZYY22-704 (AHUB) China, Gansu
Lobo hallia semis e ilis ZYY22-715 (AHUB) China, Gansu
Lobo hallia s ipi a a XY22-887 (KUN-L) China, Xinjiang
Lobo hallia s ipi a a ZYY22-616 (KUN-L) China, Xinjiang
Lobo hallia s ipi a a ZYY22-617 (KUN-L) China, Xinjiang
Lobo hallia s ipi a a 22-72871 (KUN-L) China, Xinjiang Holo ype
Lobo hallia alphoplaca SK A20 Uk aine
Lobo hallia alphoplaca O–L–200411 No way
Lobo hallia melanaspis No din 6622 Sweden
Lobo hallia melanaspis Owe-La sson 8943a No way
Lobo hallia iqbalii LAH37149 Pakis an Holo ype
Lobo hallia iqbalii LAH37150 Pakis an
Lobo hallia pakis anica LAH37137 Pakis an Holo ype
Lobo hallia pakis anica LAH37139 Pakis an
Lobo hallia pul ina a CH-93 Pakis an Pa a ype
Lobo hallia pul ina a PN-04 Pakis an
Lobo hallia polypycnidia a XY20-808 (KUN-L) China, Qinghai
Lobo hallia polypycnidia a 20-68413 (KUN-L) China, Qinghai
Lobo hallia polypycnidia a XY20-3034 (KUN-L) China, Qinghai Holo ype
Lobo hallia polypycnidia a 20-67134 (KUN-L) China, Qinghai
Lobo hallia polypycnidia a 20-68672 (KUN-L) China, Qinghai
Lobo hallia polypycnidia a 20-67303 (KUN-L) China, Qinghai
Lobo hallia polypycnidia a 20-67160 (KUN-L) China, Qinghai
Lobo hallia polypycnidia a 20-68878 (KUN-L) China, Xizang
Lobo hallia hedinii 18-59518 (KUN-L) China, Gansu
Lobo hallia hedinii 20-68298 (KUN-L) China, Qinghai
Lobo hallia hedinii 22-71155 (KUN-L) China, Xizang
Lobo hallia hedinii 20-68147 (KUN-L) China, Qinghai
Lobo hallia benzilanensis 18-60350 (KUN-L) China, Yunnan Holo ype
Lobo hallia c assima gina a Wang 20122565 China, Inne Mongolia Holo ype
Lobo hallia c assima gina a Tong 20122583 China, Inne Mongolia
Lobo hallia epiadelpha Pauko 1881 Russia Holo ype
Lobo hallia pulch a 19-66027 (KUN-L) China, Xizang Holo ype
Lobo hallia pulch a XY20-3169 (KUN-L) China, Qinghai
‘Lobo halliahelanensis ’D.B. Tong 20122518 China, Inne Mongolia
‘Lobo hallia helanensis ’D.B. Tong 20122791 China, Inne Mongolia
Lobo hallia subdi ac a a . imosa ZYY22-647 China, Xinjiang
Lobo hallia subdi ac a a . imosa Wang e al. 22-72975 China, Xinjiang
Lobo hallia subdi ac a F olo 178-1 Russia
Lobo hallia subdi ac a ZYY22-628 (AHUB) China, Xinjiang
Lobo hallia b achyloba 22-72815 (KUN-L) China, Xinjiang
Lobo hallia b achyloba F olo 357 Russia Holo ype
Lobo hallia elobula a LAH37153 Pakis an Holo ype
Lobo hallia elobula a LAH37154 Pakis an
A os elia magni ica Da ydo 21959 Russia
A os elia magni ica Da ydo 21961 Russia
Aspiciliella cup eoglauca Sipman & Raus 61847 G eece
Aspiciliella po osan ana Sipman 63019 Po ugal
Aspiciliella in e mu ans Zake i 40503 A menia
Aspiciliella in e mu ans Palice 11394 Czech Republic
Megaspo a e ucosa Zuo NJ2018074 An a c ica
Megaspo a e ucosa Sipman 55434 I an
Megaspo a imiso edia a Sp ibille WP642 Canada
Ci cina ia gibbosa No din 5878 Sweden
Ci cina ia caesiocine ea Tibell 22612 Sweden
Ci cina ia lep osescens No din 5906 Sweden
Ci cina ia a ida Wheele 5819 USA
Ci cina ia ho mannii Wheele 3634 USA
Ci cina ia esculen a Owe-La sson 9796 Russia
Sagedia mas uca a Wheele 6467 No way
Sagedia zona a Owe-La sson 8942 No way
Sagedia simoensis Owe-La sson 9000 No way
Sagedia simoensis Wheele 6288 No way
Oxne ia ia haey enii No din 5997 Sweden
Oxne ia ia mashiginensis No din 5790 Sweden
Oxne ia ia supe egens No din 6023 Sweden
Oxne ia ia pe mu a a No din 6027 Sweden
Aspicilia cine ea Wheele 6277 No way
Aspicilia indissimilis No din 5943 Sweden
Aspicilia epiglyp a No din 6303 Sweden
Aspicilia sub adians Wheele 5070 USA
Aspicilia boykinii Wheele 7263 USA
Aspicilia angelica Wheele 3535 USA
Aspicilia oli aceob unnea Owe-La sson 8771 USA
Aspicilia umosa Wheele 6802 USA
32/0.44
46/0.83
96/1.00
24/0.61
100/1.00
100/1.00
98/1.00
69/0.75
100/1.00
100/1.00
100/1.00
65/0.99
96/1.00
86/1.00
78/0.74
100/1.00
100/1.00
98/1.00
97/0.90
40/0.58
26/0.45
48/0.94
100/1.00
100/1.00
52/0.80
100/1.00
76/1.00
98/1.00
94/1.00
53/0.90
61/0.95
86/0.98
86/1.00
100/1.00
97/1.00
47/0.57
47/0.74
89/1.00
84/0.63
60/1.00
44/0.52
100/1.00
100/1.00
85/1.00
33/0.42
100/1.00
55/0.58
100/1.00
87/1.00
100/1.00
100/1.00
59/0.90
100/1.00
100/1.00
75/0.78
100/1.00
54/0.59
100/1.00
100/0.99
98/1.00
46/0.55
86/1.00
39/0.98
100/1.00
49/0.98
100/1.00
100/1.00
100/1.00
100/1.00
80/1.00
100/1.00
100/1.00
100/1.00
100/1.00
100/1.00
82/1.00
27/0.22
33/0.90
45/1.00
92/1.00
100/1.00
30/0.49
98/1.00
100/1.00
86/1.00
92/1.00
99/1.00
100/1.00
45/0.93
34/0.51
53/0.34
81/1.00
99/1.00
99/1.00
66/0.95
100/1.00
100/1.00
80/0.98
98/1.00
100/1.00
64/0.40
86/1.00
99/0.88
23/0.39
99/1.00
56/0.98
100/1.00
100/1.00
100/1.00
64/0.94
87/0.94
100/1.00
100/1.00
84/1.00
70/0.99
38/0.51
100/1.00
100/1.00
66/0.81
59/1.00
T ee scale: 0.05
Lobo hallia
Megaspo aceae
A os elia
Aspiciliella
Megaspo a
Ci cina ia
Sagedia
Oxne ia ia
Aspicilia
Teu oa
An idea
Aspilidea
Figu e 1. Bayesian phylogene ic ee o Lobo hallia in e ed om he conca ena ed alignmen o n ITS-n LSU-m SSU se-
quences. Maximum likelihood boo s ap alues (MLBP) and Bayesian pos e io p obabili ies (BPP) a e indica ed a he
nodes. Thickened b anches indica e MLBP ≥ 70% and BPP ≥ 0.95. Newly desc ibed species a e highligh ed in ed, newly
eco ded species in China in blue, and newly gene a ed sequences in bold.
213
MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554
Lun Wang e al.: An o e iew o he genus Lobo hallia in China
S a is ical suppo alues a e labeled on he b anch when he ML boo s ap
p opo ion (MLBP) ≥ 70% and Bayesian pos e io p obabili ies (BPP) ≥ 0.95.
The phylogene ic analysis esol ed 33 species o Lobo hallia in o h ee
majo clades (Clade I–III). Clade I (MLBP/BPP ≥96/1.00) ep esen s he co e
g oup o Lobo hallia, exhibi ing pheno ypic di e si y spanning he genus. The
basal lineage o his clade is Lobo hallia b achyloba, cha ac e ized by i s sho
ma ginal lobes, 1–7 apo hecia pe a eole, and c yp olecano ine apo hecia
(Pauko e al. 2019). Mos species wi hin Clade I o med highly suppo ed
monophyle ic g oups, wi h he excep ion o L. epiadelpha and L. benzilanen-
sis, which emained as single ons. Some backbone nodes, howe e , ecei ed
only weak suppo , po en ially indica ing he insu icien in o ma i e si es o
he sequences. Wi hin Clade I, eigh no el lineages we e iden i ied: se en de-
sc ibed as new species, one con i med as he species L. hedinii ha was o igi-
nally desc ibed om Gansu P o ince, China, o which no sequence was a ail-
able be o e his esea ch. Lobo hallia pulch a and L. c assima gina a clus e ed
oge he wi h weak suppo ; bo h species con ain s ic ic acid bu di e in hal-
lus colo and apo hecial ype. Ano he new species, Lobo hallia s ipi a a, also
con ains s ic ic acid bu is dis inguished by i s pale b own o b own hallus,
and i s dis inc p uinose apo hecial disc wi h a b own halline ma gin. Lobo h-
allia polypycnidia a and L. hedinii o med sis e lineages wi h s ong suppo
(87/1.00). Al hough se e al sho -b anched, highly suppo ed subclades we e
obse ed wi hin L. polypycnidia a, we ea ed hem as he same species due
o hei consis en mo phology and o e lapping geog aphical dis ibu ions,
ollowing he species delimi a ion amewo k o Ca s ens e al. (2013). Lo-
bo hallia ub a and L. complana a clus e ed oge he wi h weak suppo bu
a e mo phologically dis inc , di e ing in hallus colo and hickness (whi e
and hick s. o angish g ay and hin) and he p esence e sus absence o
gy opho ic acid. Lobo hallia pseudop uinosa, L. p uinosa, and L. densip uinosa
ga he ed in a subclade wi h high suppo (100/1.00).
Clade II consis ed solely o Lobo hallia elobula a. This clade is dis inguished
om Clades I and III by he p esence o p o hallus, a cha ac e is ic commonly ound
in he genus Aspicilia wi hin Megaspo aceae. Clade III (76/1.00) was esol ed
as a monophyle ic lineage wi h Lobo hallia wangii a he base. Wi hin his clade,
Lobo hallia hyd ocha is, L. lobula a, L. adiosa, and L. ecedens o med a s ongly
suppo ed subclade (100/1.00), in which all species consis en ly displayed g ay o
da k g ay halli. The emaining species, Lobo hallia ce noho skyana, L. che esina,
L. c enula a, L. de e mina a, and L. pel as ic oides, o med a dis inc , well-suppo ed
subclade (70/0.99) cha ac e ized by whi e o o -whi e halli.
Taxonomy
New species
Lobo hallia benzilanensis Lun Wang & Y. Y. Zhang, sp. no .
Fungal Names: FN 572942
Fig. 2
E ymology. The speci ic epi he e e s o Benzilan Town, he ype locali y o
his species.
220
MycoKeys 125: 205–244 (2025), DOI: 10.3897/mycokeys.125.173554
Lun Wang e al.: An o e iew o he genus Lobo hallia in China
Xin-Yu Wang e al. XY19-2271 (KUN-L 73043) • Chamdo Ci., Jiangda Co., Tong-
pu Vil., 31°38'30.46"N, 98°26'11.62"E, al . 3948 m, on limes one, 23 Sep embe
2020, Li-Song Wang e al. 20-68878 (KUN-L 77062), 20-68840 (KUN-L 77024).
Lobo hallia pseudop uinosa Lun Wang & Y. Y. Zhang, sp. no .
Fungal Names: FN 572945
Fig. 5
E ymology. The epi he indica es ha his species is simila o Lobo hallia p u-
inosa in mo phology.
Diagnosis. This species is cha ac e ized by i s igh ly adna e and hin hallus,
a eola e cen e and sho ly loba e ma gin; plane, whi e o b ownish g ay uppe
su ace wi h ine g anula p uina; c yp olecano ine apo hecia; la ge ascospo es
(10.0-)11.8–13.3–14.8(-17.0) × (8.0-)8.5–9.3–10.0(-11.0) µm; and i s long co-
nidia (5–)6–8(–9) × 1 µm.
Holo ype. China • Xizang Au onomous Region: Shiga se Ci., Ding i Co.,
Zhaguo Vil., 28°35'23.26"N, 86°53'55.21"E, al . 4320 m, on ock, 27 July 2019,
Li-Song Wang e al. 19-64075 (KUN-L 68571).
Desc ip ion. Thallus igh ly adna e o he subs a e, up o 3 cm ac oss, cen-
ally a eola e, 1–2 mm hick, ma ginally loba e, 0.3–0.8 mm hick. A eoles
(0.3–)0.6–1(–1.5) mm wide, angula o ounded, la , no cons ic ed a base;
in e spaces be ween a eoles 0.05–0.1 mm wide. Lobes adia e, sho , plane,
apices equal o sligh ly wide han base, 1–2 mm long, base 0.5–1 mm wide,
apex 0.5–1.5 mm wide. Uppe su ace whi e o pale b ownish g ay, p uinose,
b ownish whe e p uina hin. Uppe co ex pa aplec enchyma ous, 30–50 µm
hick, con aining da k b own g anules (soluble in K); epinec al laye wi h dense
g ayish-black g anules (insoluble in K), 10–40(–50) µm hick. Algal laye 70–
100(–150) µm hick, discon inuous; pho obion chlo ococcoid, (6–)8–20 µm
diame e . Medulla 0.1–0.5 mm hick, opaque, illed wi h g ayish-black g anules
(insoluble o pa ially soluble in K). Lowe co ex absen .
Apo hecia c yp olecano ine, common, g ouped in he cen e , 1–3 pe a eole,
angula , (0.1–)0.3–0.8(–1.0) mm in diame e ; disc conca e, b ownish o black,
ma , p uinose; apo hecial ma gin indis inc , concolo ous wi h he hallus. Ex-
ciple na ow, widening o 10–30(–50) μm in he uppe mos pa . Epi hecium,
hymenium and subhymenial laye s combined (100–)125–150(–175) µm high;
epi hecium 5–15 µm hick, gela inous; epihymenium 15–25 µm high, wi h da k
b own o b ownish g anules (soluble in K), N–; hymenium (70–)90–100(–110)
µm high, hyaline, I+ blue; subhymenial laye s 50–75 µm high, hyaline, I+ blue; al-
gal laye ela i ely con inuous below hypo hecium, 50–70 μm high; pa aphyses
submonili o m, simple, sep a e, wi h 1–3 uppe mos cells sho e and wide
han he basal cells, 4–6 μm wide (basal cells ca. 2 μm wide); asci cla a e,
Aspicilia- ype, hyaline, 8-spo ed, 60–75 × 15–25 µm; ascospo es simple, hy-
aline, b oadly ellipsoid, a ely sphe ical, (10.0-)11.8–13.3–14.8(-17.0) × (8.0-
)8.5–9.3–10.0(-11.0) µm (n = 86), l/w a io (1.0-)1.2–1.4–1.6(-2.1), wall ca.1
µm hick. Pycnidia a e, punc i o m, sligh ly con ex, 0.05–0.1 mm diame e ,
os iole b own o da k b own, conidia bacilli o m, hyaline, (5–)6–8(–9) × 1 µm.
Chemis y. Co ex K+ pale yellow o o ange, P–. Medulla K+ yellow o o -
ange- ed, P+ yellow, C–, KC–. Con aining no s ic ic and conno s ic ic acids.
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Habi a and dis ibu ion. Saxicolous. Cu en ly known only in Qinghai, Xizang
P o ince, China.
No es. Specimens o his new species we e collec ed om Qinghai and Xi-
zang P o inces, China, a ele a ions anging om 3000 o 4500 m. The o e -
all mo phology o hese specimens is uni o m, wi h he excep ion o ouche
20-67831 (KUN-L 76010), which exhibi s a hinne uppe co ex composed o
loosely a anged hyphae.
Phylogene ically, Lobo hallia p uinosa and L. densip uinosa a e he closes
ela i es o he new species. L. p uinosa can be dis inguished by i s lecano ine
apo hecia a ma u i y and b oade lobes. L. densip uinosa di e s in i s whi ish o
ligh g eenish g ay and en i ely p uinose hallus, apo hecial disc da k oli aceous
when we and blackish when d y, and he absence o conno s ic ic acid (Kou e al.
2013; Ash a e al. 2022). Lobo hallia pakis anica esembles L. pseudop uinosa in
i s whi e, loba e hallus bu di e s in possessing la ge cen al a eoles, b oade
ma ginal lobes, and lacking o seconda y me aboli es (Zul iqa e al. 2022).
Wi hin Lobo hallia, L. b achyloba, L. pla yca pa L. pul ina a and his new species
sha e an a eola e hallus wi h loba e ma gins, a whi ish o whi ish-g ay uppe su -
ace, and he p esence o no s ic ic acid. Ne e heless, each can be dis inguished
Figu e 5. Lobo hallia pseudop uinosa (KUN-L 68571, holo ype). A. Thallus; B. A eoles and apo hecia; C. Lobes; D. Sec ion
o hallus; E. Sec ion o hallus (LCB); F. Ve ical sec ion o apo hecium; G. Ve ical sec ion o apo hecium (Lugol’s solu-
ion); H. Ve ical sec ion o apo hecium (LCB); I. Ascus and ascospo es; J. Pa aphyses (LCB); K. Conidia. Scale ba s:1
mm (A); 0.2 mm (B, C); 50 µm (D, F); 20 µm (E, G, H); 5 µm (I, J, K).
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by speci ic cha ac e s: L. b achyloba has spa se halline and 1–7 apo hecia pe
a eole (Pauko e al. 2019); L. pla yca pa possesses la ge a eoles, ewe and
non-agg ega ed apo hecia, and limi ed dis ibu ion in No h A ica (Alge ia) (Zu-
l iqa e al. 2022); And L. pul ina a exhibi s a hinne hallus and uppe co ex
(10–15 µm), and lecano ine apo hecia (Zul iqa e al. 2023). Addi ionally, Lobo h-
allia che esina, L. con o e sa, and L. lac eola also ha e a igh ly adna e, whi ish
o whi ish–g ay hallus, bu hese species di e om L. pseudop uinosa in hei
absen o indis inc ma ginal lobes and hei di e en seconda y me aboli es.
Addi ional specimens examined. Lobo hallia pseudop uinosa. China • Xi-
zang: Shiga se Ci., Ding i Co., Zhaguo Vil., G219, 28°35'09.99"N, 87°03'42.35"E,
al . 4306 m, on su ace o wea he ed shale in open a ea, 16 June 2022, Li-
Song Wang e al. 22-71222 (KUN-L 85846), Yan-Yun Zhang ZYY22-324
(KUN-L 81905), ZYY22-326 (KUN-L 81907) • Ang en Co., 29°19'01.32"N,
87°01'59.60"E, al . 4524 m, on ock, 19 July 2019, Li-Song Wang e al. 19-
63621 (KUN-L 68115) • Qinghai P o .: Yushu Tibe an Au onomous P e ec-
u e, Qumalai Co., Qumahe Vil., 34°54'50.67"N, 94°46'20.74"E, al . 4396 m, on
limes one, 17 Sep embe 2020, Li-Song Wang e al. 20-68331 (KUN-L 76512)
• Zeku Co., Maixiu Town, 35°15'53.57"N, 101°52'30.23"E, al . 3143–3163 m,
on ock, 8 July 2022, Xin-Yu Wang and Min Ai XY22-1089 (KUN-L 84907),
XY22-1088 (KUN-L 84906), An-Cheng Yin and Han-Xiang Chen 22-72438
(KUN-L 87064) • Jiuzhi Co., NianBaoYuZe Na ional Geopa k, 33°13'57.57"N,
100°55'58.56"E, al . 4063 m, on sandy soil, 8 Sep embe 2020, Li-Song Wang
e al. 20-67831 (KUN-L 76010).
Lobo hallia p uinosa. China • Nei Mongol: Bayanho Town, Helan Moun-
ain, al . 1600 m, on ock, 17 Augus 2011, H.Y.Wang 20123575, 20123154,
20123282, 20123626 (SDNU) • U a Rea Banne Co., al . 1600 m, on ock, 19
Augus 2011, H.Y.Wang 20122917 (SDNU), D.B.Tong 20123276, 20123888
(SDNU) • Qinghai P o .: Hualong Hui Eh nic Au onomous Co., Yashiga Town,
36°03'52.78"N, 101°57'03.44"E, al . 2002 m, on ock, 8 July 2022, Li-Song Wang
e al. 22-73170 (KUN-L 87648).
Lobo hallia pulch a Lun Wang & Y. Y. Zhang, sp. no .
Fungal Names: FN 572946
Fig. 6
E ymology. The epi he e e s o he beau i ul mo phology o he hallus o his
species.
Diagnosis. Thallus a eola e wi h adia e lobes, uppe su ace whi e and p u-
inose; apo hecia lecano ine, adna e and ounded, 1–2 apo hecia pe a eole,
apo hecial ma gin dis inc and pe sis en , disc plane o sligh ly con ex, no o
a ely wi h ain p uina, con aining no s ic ic and s ic ic acids.
Holo ype. China • Xizang Au onomous Region: Shiga se Ci., Ding i Co.,
Zhaguo Vil., 28°35'24.49"N, 86°53'58.22"E, ele . 4293 m, on ock, 27 July 2019,
Li-Song Wang e al. 19-66027 (KUN-L 70435).
Desc ip ion. Thallus igh ly adna e o he subs a e, 0.3–2 mm hick, cen-
ally a eola e, ma ginally loba e. A eoles 0.5–1.2 mm wide, angula o ound-
ed, no cons ic ed a base, in e spaces be ween a eoles 0.05–0.1 mm wide.
Lobes adia e, sho , 1–2.5 mm long, base wid h (0.3–)0.6–0.8(–1) mm, apex
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wid h 0.6–2 mm. Uppe su ace mode a ely con ex, whi e, pa ially pale b own
on ma ginal lobes, p uinose. Uppe co ex pa aplec enchyma ous, e en, 30–
40 μm hick, con aining b own g anules (soluble in K); epinec al laye gela i-
nous, 10–20 μm hick. Algal laye 125–150 μm high, discon inuous; pho obion
chlo ococcoid, cells 6–18 µm diame e . Medulla 0.3–0.6 mm hick, opaque,
illed wi h g ay-black g anules (insoluble in K). Lowe co ex absen .
Apo hecia lecano ine, common, 1–2 pe a eole, simple o g ouped in he
cen e , o bicula o sligh ly angula by p essu e om adjacen apo hecia, (0.3–
)0.6–2 mm diame e , ini ially imme sed, la e sessile, no o sligh ly cons ic ed
a base; disc plane, ma , b ownish-black o black, ep uinose o ain ly p uinose,
shallowly issu ed when o e ma u e; apo hecial ma gin en i e, concolo ous wi h
hallus, well-de eloped and pe sis en , (0.05–)0.1–0.2(–0.25) mm. Exciple na -
ow, widening o 70–80 μm in he uppe mos pa . Epi hecium, hymenium and
subhymenial laye s combined 125–160 µm high; epi hecium 5–15 µm high, epi-
hymenium 12.5–20 μm high, illed wi h da k b own o b own g anules (mos ly
soluble in K), N–; hymenium 90–100 μm high, hyaline, I+ da k blue; subhymenial
laye s ca. 50 μm high, hyaline, I+ da k blue; discon inuous algal laye benea h
hypo hecium; pa aphyses simple, sep a e, submonili o m a he ips, 1–4 uppe -
mos cells sho e and b oade han basal cells, 3–5 μm wide (basal cells 2–3 μm
Figu e 6. Lobo hallia pulch a (KUN-L 70435, holo ype). A. Habi a ; B. Thallus; C. Lobes; D. Apo hecia; E. Ve ical sec ion
o apo hecium; F, G. Sec ion o hallus (G in LCB); H. Ve ical sec ion o apo hecium (LCB); I. Ascus; J. Pa aphyses (LCB);
K. Ascospo es; L. Conidia (LCB). Scale ba s: 1mm (B); 0.5 mm (C); 0.2 mm (D); 100 µm (E, F, G); 50 µm (H); 5 µm (I, J, K, L).
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wide); asci cla a e, Aspicilia- ype, hyaline, 8-spo ed, 70–80 × 20–30 µm; asco-
spo es simple, hyaline, ellipsoid o b oadly ellipsoid, (10.0-)10.8–12.3–13.7(-
16.0) × (8.0-)7.8–8.5–9.1(-10.0) µm (n = 30), l/w a io (1.1-)1.3–1.5–1.7(-1.9),
wall ca.1 µm hick. Pycnidia ew, punc i o m, la o sligh ly con ex, 0.15–0.2 mm
diame e ; os iole b ownish black; conidia bacilli o m, hyaline, 6–8 × 1 µm.
Chemis y. Co ex K+ pale yellow o K–, P–; medulla K+ yellow o o ange- ed,
P+ yellow o o ange- ed, C–, KC– o KC+ pale yellow; no s ic ic and s ic ic acids
de ec ed by TLC, some specimens addi ionally con ain c yp os ic ic, conno s -
ic ic and cons ic ic acids.
Habi a and dis ibu ion. Saxicolous. Cu en ly known om Xizang and Qin-
ghai P o inces, China.
No es. Lobo hallia pseudop uinosa and L. p uinosa esemble L. pulch a in
ha ing an a eola e hallus wi h adia e lobes and a whi e p uinose su ace, bu
di e in hei indis inc apo hecial ma gin, dis inc ly p uinose discs, and he
absence o s ic ic acid (Kou e al. 2013). Lobo hallia wangii di e s om L. pul-
ch a in i s p edominan ly c yp olecano ine apo hecia, N+ epihymenium, and he
absence o bo h ma ginal lobes and s ic ic acid. Lobo hallia pakis anica and
L. pul ina a also ha e a whi e hallus wi h cen al a eoles and ma ginal lobes,
bu di e om L. pulch a in hei hinne halli, apo hecia ha a e no cons ic -
ed a base, and in he lack o s ic ic acid (Zul iqa e al. 2022, 2023).
Addi ional specimens examined. China • Xizang Au onomous Region: Tin-
g i Co., Zhaguo Vi., 28°35'24.49"N, 86°53'58.22"E, al . 4256–4299 m, on ock,
27 July 2019, Li-Song Wang e al. 19-65686 (KUN-L 70194), 19-65652 (KUN-L
70159) • Dagze Co., Bangdui Vi, 29°44'06.51"N, 91°24'55.29"E, al . 3700 m, on
ock, 16 July 2019, Li-Song Wang e al. 19-64597 (KUN-L 69093) • Qinghai: Yushu
Tibe an Au onomous P e ec u e, Zaduo Co., 32°52'43.64"N, 95°20'29.61"E, al .
4074 m, on ock, 20 Sep embe 2020, Li-Song Wang e al. 20-67281 (KUN-L
75458) • Ado Vi., 32°52'43.35"N, 95°20'30.19"E, al . 4080 m, on ock, 20 Sep-
embe 2020, Xin-Yu Wang e al. XY20-3169 (KUN-L 79525).
Lobo hallia ub a Lun Wang & Y. Y. Zhang, sp. no .
Fungal Names: FN 572947
Fig. 7
E ymology. The speci ic epi he e e s o he ed eac ion o medulla in C spo - es .
Diagnosis. Thallus a eola e wi h a loba e ma gin, uppe su ace whi e wi h
pale b own inge, p uinose; a eoles aised when bea ing apo hecia o pycnidia;
apo hecia lecano ine, disc p uinose; pycnidia p o uding; medulla C+ ose– ed,
con ain gy opho ic acid.
Holo ype. China • Xinjiang Uygu Au onomous Region: Hejing Co., Balun ai
Vil., 42°51'56.45"N, 86°27'09.16"E, al . 2026 m, on ock, 02 July 2022, Xin-Yu
Wang and Min Ai XY22-897 (holo ype: KUN-L 84715).
Desc ip ion. Thallus closely adna e o subs a e, up o 5 cm ac oss, a eo-
la e, 1–4 mm hick, ma ginally loba e, 0.4–1 mm hick. A eoles (0.5–)0.8–2(–
2.4) mm wide, con iguous, angula o ounded, non-cons ic ed a base, in e -
spaces be ween a eoles 0.1–0.2 mm wide. Lobes sligh ly b oade in he apex,
2–3 mm long, 0.7–1.4 mm wide a base, 0.6–2(–2.5) mm wide a apex. Uppe
su ace con ex, c acked, p uinose, whi e wi h pale b own inge whe e p uina is
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hin. Uppe co ex pa aplec enchyma ous, 20–40 μm hick, con aining b own
g anules (soluble in K); epinec al laye composed o coa se pla e-like c ys als,
gela inous ma e ial, and g ay-black g anules (insoluble in K), 10–30 μm hick. Al-
gal laye 75–150 μm high, discon inuous, in e up ed by hyphae, 10–60 μm wide;
pho obion chlo ococcoid, cells 8–20 µm diame e . Medulla 0.5–1 mm high,
illed wi h g ay–black g anules (gene ally insoluble in K). Lowe co ex absen .
Apo hecia lecano ine, common, usually 1 pe a eole, (0.3–)0.6–1.5(–
2.0) mm diame e , ounded, sessile, sligh ly cons ic ed a base when ma u e;
disc sligh ly conca e o plane, black, p uinose; apo hecial ma gin en i e, pe -
sis en , 0.1–0.25 mm wide, p uinose, concolo ous wi h hallus. Exciple na ow,
Figu e 7. Lobo hallia ub a (KUN-L 84715, holo ype); A. Thallus; B. Apo hecia; C. Pycnidia; D. Lobes; E. Sec ion o hallus
(LCB); F. Sec ion o hallus (in K); G. Ve ical sec ion o apo hecium; H. Ve ical sec ion o apo hecium (LCB); I, J. Ve ical
sec ion o pycnidium; K. Ascus and ascospo es; L. Pa aphyses (le in LCB, igh in K); M. Conidia (LCB). Scale ba s: 2mm
(A); 0.5 mm (D); 0.2 mm (B, C); 50 µm (G, I, J); 20 µm (E, F, H); 10 µm (K, M); 5 µm (L).
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up o 40 μm wide. Epi hecium, hymenium and subhymenial laye s combined
(150–)175–225(–250) µm high; epi hecium 5–20 µm high; epihymenium
20–50 μm high, illed wi h da k b own g anules (soluble in K), N–; hymenium
100–150 μm high, hyaline, I– o I+ weakly bluish in lowe pa ; subhymenial
laye s 50–100 μm high, hypo hecium 40–50 μm, hyaline, I+ blue; algal laye be-
low hypo hecium discon inuous, 50–60 μm high; pa aphyses simple, sep a e,
submonili o m a he ips, wi h 2–3 uppe mos cells sho e and b oade han
basal cells, 3–6 μm wide (basal cells 2–3 μm wide); asci cla a e, Aspicilia- ype,
hyaline, 8-spo ed, 70–85 × 20–30 µm; ascospo es simple, hyaline, b oadly ellip-
soid, (10.0-)11.9–13.1–14.2(-15.0) × (8.0-)8.7–9.4–10.0(-11.0) µm (n = 48), l/w
a io (1.0-)1.3–1.4–1.5(-1.8), wall ca.1 µm hick. Pycnidia common, p o uding,
0.1–0.25 mm in diame e , some imes elonga ed, 0.5(–0.7) × 0.2 mm (leng h ×
wid h); os iole b ownish o da k b own; conidia bacilli o m, hyaline, 5–6 × 1 µm.
Chemis y. Co ex: K+ pale yellow, P–; medulla: K+ pale yellow o o ange o
K–, P–, C+ ose– ed, KC+ ed o pale yellow; con aining gy opho ic, no s ic ic,
s ic ic, conno s ic ic and cons ic ic acids.
Habi a and dis ibu ion. Saxicolous. Cu en ly known only om Xinjiang, China.
No es. This species is eadily ecognized by i s C+ ose– ed medulla y eac-
ion, esul ing om he p oduc ion o gy opho ic acid. Lobo hallia semis e ilis
esembles his species in i s whi e o g ay hallus and p o uding pycnidia, bu
di e s in being e icolous and lacking gy opho ic and s ic ic acids (Zhang e
al. 2020). Lobo hallia p uinosa sha es he cha ac e is ic whi e p uina on bo h
hallus and discs, bu di e s in i s hinne hallus, punc i o m pycnidia, and he
p esence o no s ic ic and conno s ic ic acids as he majo seconda y me ab-
oli e (Kou e al. 2013). Lobo hallia pulch a is simila o L. ub a in possessing
whi e hallus and lecano ine apo hecia, bu di e s in ha ing ep uinose discs
and lacking gy opho ic acid.
Addi ional specimens examined. China • Xinjiang Uygu Au onomous Region:
Hejing Co., Balun ai Town, 42°51'56.45"N, 86°27'09.16"E, al . 2026–2046 m, on
ock, 2 July 2022, An-Cheng Yin and Han-Xiang Chen 22-72301 (KUN-L 86927) •
Na ional Highway 216, 42°51'56.39"N, 86°27'09.31"E, al . 2100 m, on limes one,
2 July 2022, Li-Song Wang e al. 22-72886 (KUN-L 87373).
Lobo hallia s ipi a a Lun Wang & Y. Y. Zhang, sp. no .
Fungal Names: FN 572948
Fig. 8
E ymology. The epi he e e s o he a eoles wi h a cons ic ed and s ipi a e base.
Diagnosis. Thallus b own, hick, cen ally a eola e, a eoles wi h a s ipi a e
base, ma ginally loba e, ma gin o lobes ee om he subs a e wi h g anula
p uina on he uppe su ace; apo hecia lecano ine, cons ic ed a he base, disc
wi h conspicuous whi e p uina, apo hecial ma gin pe sis en , ep uinose, b own
o o ange-b own; hymenium pale b own.
Holo ype. China • Xinjiang Uygu Au onomous Region: Hejing Co., along
oad G218, 42°54'22.90"N, 86°17'36.95"E, al . 2228 m, on ock, 01 July 2022,
Li-Song Wang e al. 22-72871 (KUN-L 87358).
Desc ip ion. Thallus ela i ely loosely a ached o he subs a e, up o 4–6 cm
ac oss, cen ally a eola e, 3–6(–8) mm hick, ma ginally loba e, (0.5–)1–2 mm
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hick, ee om he subs a e. A eoles angula (0.6–1.2 mm wide) o elonga ed
(1.5–2 × 0.6–1 mm wide), base cons ic ed in o a s ipe, up o 4 mm long, in e spac-
es be ween a eoles 0.1–0.3 mm wide. Lobes long, simple o 2–3 b anched, apices
sligh ly wide han base, 3–5 mm long, 0.6–1.5 mm wide a base, 0.6–2.5 mm
wide a apex. Uppe su ace la o sligh ly con ex, ma , b own, hinly p uinose,
g anula p uina p esen on he ma gin o lobes. Uppe co ex pa aplec enchyma-
ous, e en, 40–50 μm hick, inspe sed wi h b ownish and g ay g anules (soluble in
K); epinec al laye gela inous, 10–15 μm hick. Algal laye 75–150 μm high, discon-
inuous; pho obion chlo ococcoid, cells 10–25 µm diame e . Medulla 0.6–3 mm
high, illed wi h g ay-black g anules (gene ally insoluble in K). Lowe co ex absen .
Figu e 8. Lobo hallia s ipi a a (KUN-L 87358, holo ype). A. Thallus; B. P o ile o hallus; C. S ipe; D. Lobes; E. Apo hecia;
F. Pycnidium; G, H. Sec ion o hallus (H in K eagen ); I. Ve ical sec ion o apo hecium (LCB); J. Ve ical sec ion o apo h-
ecium; K. Uppe co ex and epinec al laye (LCB); L. Ascus; M. Pa aphyses (LCB); N. Ascospo es; O. Conidia. Scale ba s:
2 mm (A); 1 mm (B, C); 0.5 mm (D); 0.2 mm (E, F); 100 µm (G, H); 50 µm (I, J); 20 µm (K); 10 µm (L, M, N, O).
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Apo hecia lecano ine, common, 1–2 pe a eole, simple o g ouped, o bicu-
la o angula by p essu e, (0.3–)0.5–1.5(–1.8) mm wide, cons ic ed a base
in ma u i y; disc conca e o plane, b own o black, conspicuously p uinose;
apo hecial ma gin en i e, pe sis en , ep uinose, b own o o ange-b own, 0.1–
0.25(–0.3) mm wide. Exciple na ow, widening o 60–80 μm in he uppe mos
pa . Epi hecium, hymenium and subhymenial laye s combined 125–175 µm
high; epi hecium 5–15 µm high, wi h hyaline pla e-like c ys als; epihymenium
12.5–37.5 μm high, illed wi h da k b own g anules (soluble in K), N–; hyme-
nium 100–120 μm high, pale b own (soluble in K), I+ da k blue; subhymenial
laye s 50–75 μm high, hypo hecium 40–50 μm, pale b own (soluble in K), I+
da k blue; algal laye below hypo hecium spa se o absen ; pa aphyses simple,
sep a e, submonili o m a he ips, wi h 2–4 uppe mos cells sho e and b oad-
e han basal cells, 3–5 μm wide (basal cells ca. 2 μm wide); asci cla a e, As-
picilia- ype, hyaline, 8-spo ed, 70–85 × 18–25 µm; ascospo es simple, hyaline,
b oadly ellipsoid o ellipsoid, (9.0-)9.8–11.4–12.9(-15.0) × (7.0-)6.8–7.6–8.4(-
11.0) µm (n = 30), l/w a io (1.0-)1.3–1.5–1.7(-2.0), wall ca.1 µm hick. Pycnidia
punc i o m, plane o sligh ly con ex, 0.1–0.25 mm diame e ; os iole b own o
da k b own; conidia bacilli o m, hyaline, 4–6 × 1 µm.
Chemis y. Co ex K+ pale yellow, P–; medulla K+ yellow, P+ o ange, C–, KC–;
con aining no s ic ic, s ic ic, c yp os ic ic and conno s ic ic acids.
Habi a and dis ibu ion. Saxicolous. Cu en ly only known om Xinjiang
P o ince, China.
No es. Lobo hallia p ae adiosa esembles L. s ipi a a in ha ing an a eola e hal-
lus wi h adia ing lobes, a cons ic ed apo hecial base, o ange-b own hallus and
apo hecial ma gins, and con aining no s ic ic acid. Howe e , he o me di e s in i s
ep uinose disc, hyaline hymenium, and lack o s ic ic acid (Kou e al. 2013; Pauko
e al. 2019; Ryan 2004). Lobo hallia semis e ilis sha es na owly elonga ed lobes
wi h g anula p uina on he ma gins, bu di e s in i s e icolous habi a , whi e o
g ay hallus, apo hecia-like pycnidia, and absence o s ic ic acid (Zhang e al. 2020).
Addi ional specimens examined. China • Xinjiang Uygu Au onomous Re-
gion: Hejing Co., Na ional Highway 218, 42°54'22.90"N, 86°17'36.95"E, al .
2228 m, on limes one, 1 July 2022, Yan-Yun Zhang ZYY22-616 (AHUB 00470),
ZYY22-617 (KUN-L 82191) • Balun ai Town, 42°54'36.56"N, 86°17'02.47"E, al .
2171 m, on ock, 1 July 2022, Xin-Yu Wang and Min Ai XY22-887 (KUN-L 84705).
Lobo hallia wangii Lun Wang & Y. Y. Zhang, sp. no .
Fungal Names: FN 572949
Fig. 9
E ymology. The epi he e e s o he Chinese lichenologis Li-Song Wang, who
collec ed he ype specimen.
Diagnosis. Thallus a eola e wi hou ma ginal lobes, uppe su ace whi e,
la ; apo hecia common, c yp olecano ine o lecano ine, disc black, p uinose,
sligh ly conca e o plane, o en shallowly issu ed a ma u i y; epihymenium N+
weakly g eenish o g een, con aining no s ic ic and conno s ic ic acids.
Holo ype. China • Xizang Au onomous Region: Shiga se Ci., Ding i Co., along
oad G219, 28°35'10.18"N, 87°03'43.62"E, al . 4311 m, on ock, 06 June 2022,
Li-Song Wang e al. 22-71215 (KUN-L 85839).
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Desc ip ion. Thallus igh ly adna e o he subs a e, 1–4 mm hick, a eo-
la e wi hou ma ginal lobes, sp eading in he ield up o 25 cm ac oss, he
collec ed specimen is agmen ed, up o 4 cm ac oss. Cen al a eoles com-
monly ounded, (0.2–)0.5–1(–1.6) mm diame e , no cons ic ed a base,
in e spaces be ween a eoles 0.1–0.2 mm wide; ma ginal a eoles ounded
o sligh ly elonga ed, 0.8–1.2 × 0.5–1.2 mm. Uppe su ace la , whi e and
p uinose. Uppe co ex pa aplec enchyma ous, 30–50 μm hick, inspe sed
wi h pale g ay g anules (soluble in K), uppe mos pa illed wi h da k b own
g anules (pa ly soluble in K), 20–30 μm hick; epinec al laye g ay da k, 20–
50 μm hick. Algal laye 75–100 μm hick, mos ly con inuous, occasionally
Figu e 9. Lobo hallia wangii (KUN-L 85839, holo ype). A. Habi ; B, C. Apo hecia; D. Sec ion o hallus (below in K eagen );
E. Uppe co ex and epinec al laye (LCB); F. Sec ion o apo hecium; G. Sec ion o apo hecium (in N eagen ); H. Sec ion
o pycnidium; I. Ascus and ascospo es; J. Pa aphyses (LCB); K. Conidia. Scale ba s: 1 mm (A); 0.2 mm (B, C); 100 µm (D,
G); 20 µm (E, F, H); 10 µm (I); 5 µm (J, K).
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g ay o o ange–b own, apo hecial ma gin g ay o o angish–b own .......
...........................................................................................L. p ae adiosa
16 Cen al hallus hin, less han 2 mm; lobes plane ......................................17
– Cen al hallus hick, mo e han 2 mm; lobes sligh ly con ex ..................20
17 Apo hecia no exceeding 3 pe a eole .......................................................18
– Apo hecia commonly mo e han 3 pe a eole ........................ L. b achyloba
18 Disc black o da k b own wi h whi e p uina ...............................................19
– Disc b own wi h o ange-g ay p uina ....................................... L. complana a
19 Apo hecia c yp olecano ine, 0.3–0.8 mm, apo hecial ma gin a he same
le el wi h disc ...................................................................L. pseudop uinosa
– Apo hecia c yp olecano ine o lecano ine, 0.7–1.2 mm, apo hecial ma gin
somewha highe han he disc ................................................... L. p uinosa
20 Pycnidia spa se, punc i o m o sligh ly p o uding a ma u i y ................21
– Pycnidia dense pe a eole, spo -like o elonga e and cu ed .......................
..............................................................................................L. polypycnidia a
21 Ma ginal lobes pa allel, disc p uinose.............................................L. hedinii
– Ma ginal lobes non-pa allel, disc ep uinose ................................. L. adiosa
Discussion
Species di e si y
Thi y-nine species o Lobo hallia a e accep ed wo ldwide in his s udy. Among
hem, wen y- wo a e p esen in China, including eigh new species (Figs 2–9),
one newly eco ded species (Fig. 10), and hi een known species, en o which
a e s udied he e (Figs 11, 12) (Magnusson 1940, 1944; Mamu e al. 2010,
2012; Kou e al. 2013; Pauko e al. 2019; Wheele e al. 2024; Zhang e al. 2020,
2024a). This esea ch signi ican ly inc eases he species di e si y o Lobo hal-
lia and shows ha China is a dis ibu ion cen e o his genus, especially in he
no h, no hwes and sou hwes . The Qinghai-Tibe Pla eau egion eme ges as
a signi ican di e si y ho spo o he genus, exhibi ing a subs an ially highe
species ichness compa ed o o he a eas. Un o una ely, no specimens om
no heas e n China we e examined, bu comp ehensi e u u e sampling may
u he en ich he species di e si y o Lobo hallia in he coun y.
Mo phological and mic oscopical cha ac e s
In Megaspo aceae, Lobo hallia is cha ac e ized by ela i ely small ascospo es
ha a ely exceed 18 μm in leng h, and sho conidia, 3–10 μm long (No din e
al. 2010; Kou e al. 2013; Pauko e al. 2019; Zul iqa e al. 2022). Addi ional
mo pho-ana omical cha ac e s include: a pa aplec enchyma ous uppe co ex,
a discon inuous algal laye ; an N- o a ely N+ sligh ly g eenish epihymenium ( s.
N+ eme ald g een in o he gene a o Megaspo aceae) (Pauko e al. 2019); sim-
ple o a ely anas omosed pa aphyses wi h swollen, submonili o m o monili-
o m ips; and a subhypo hecial algal laye (Ryan 2004; Kou e al. 2013; Pauko
e al. 2019), which may be poo ly isible in some cases (Roux e al. 2016).
Al hough An idea b ucei (Owe-La ss. & A. No din) T. B. Wheele , Aspilidea my inii
(F .) Ha ellne and A. subadunans (Vain.) T. B. Wheele , J. W. McCa hy & F yday
possess ascospo es and conidia o simila size o he Lobo hallia species, hey
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a e phylogene ically dis an (Ha ellne and Tü k 2001; Owe-La sson e al. 2007;
Wheele e al. 2024). Simila ly, species o Teu oa sha e ana omical simila i ies wi h
Lobo hallia, such as pa aphyses s uc u e, ascospo e and conidia size. Howe e ,
Teu oa di e s in ha ing non-loba e halli, lacking seconda y me aboli es, and i s
co icolous o e icolous habi a (Rico e al. 2007; Soh abi e al. 2013b).
The adi ional key mo phological ea u e o de ining he genus Lobo hallia—
he p esence o ma ginal lobes—exhibi s ansi ional a ia ion ac oss he genus
(Ha ellne 1991; Ryan 2004). Al hough i is he e o e no a eliable cha ac e o
gene ic delimi a ion, i emains aluable in species-le el iden i ica ion (Pauko
e al. 2019). Cu en ly, app oxima ely one- hi d o he species in Lobo hallia lack
o exhibi indis inc lobes, iz. L. benzilanensis, L. ce noho skyana, L. chade audi-
ana, L. che esina, L. con o e sa, L. de e mina a, L. elobula a, L. gangwondoana,
L. lac eola, L. pel as ic oides, L. ecedens, L. subdi ac a and L. wangii (Magnus-
son 1940, 1944; Roux e al. 2016; Pauko e al. 2019; Kond a yuk e al. 2020; Zu-
l iqa e al. 2022; Wheele e al. 2024). The leng h o lobes is a diagnos ic cha ac-
e a species le el (Pauko e al. 2019; Zul iqa e al. 2023), anging om sho
(< 2 mm) o ela i ely long (up o 5 mm). Mos species o Lobo hallia ha e an a e-
ola e hallus (a leas in he cen e ), wi h he ollowing excep ions: L. epiadelpha,
which is cen ally squamulose; L. zog ii, which is cen ally squamulose-a eola e
Figu e 12. O he Lobo hallia species om China examined in his s udy. A. L. alphoplaca (22-72816, KUN-L);
B. L. c assima gina a (20122418B, SDNU); C. L. c enula a (ZYY22-301, KUN-L, holo ype); D. L. lobula a (ZYY22-819, KUN-L,
holo ype); E. L. p ae adiosa (22-71752, KUN-L); F. L. p uinosa (22-73170, KUN-L); G. L. semis e ilis (ZYY22-719, KUN-L);
H. L. subdi ac a (ZYY22-628, KUN-L); I. L. adiosa (22-72256, KUN-L). Scales ba s: 1 mm (B–I).
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Lun Wang e al.: An o e iew o he genus Lobo hallia in China
(Pauko e al. 2019); and L. benzilanensis and L. gangwondoana, which con ain
mo e o less dis an simple a eoles (Kond a yuk e al. 2020).
Species delimi a ion in Lobo hallia is based on he in eg a ion o mul iple mo -
phological and chemical cha ac e s. Key hallus ea u es o iden i ica ion include
hallus hickness ( a ying om < 2 mm o 8 mm), a eoles size, and he p esence
o p uina. Apo hecial ype (e.g., c yp olecano ine, c yp olecano ine o lecano ine,
lecano ine), disc colo and p uina, and halline ma gin wid h and de elopmen a e
also equen ly employed o species delimi a ion. Addi ional pheno ypic cha ac-
e s u he suppo species delimi a ion, including habi a condi ions and sub-
s a e ype; hallus a achmen , g ow h o m and diame e , he wid h and dep h o
in e -a eola c acks; he p esence o absence o lobules and o basal cons ic ion
in a eoles; smoo h o imose uppe su ace; and he co ex and medulla hickness.
Rep oduc i e ai s include he numbe o apo hecia and pycnidia pe a eole; he
p esence o absence o a cons ic ed apo hecial base; disc shape (conca e, plane,
con ex) and ex u e (shiny/ma , wi h o wi hou issu es); epihymenium eac ions
(N− o N+ ligh g een), hymenium and hypo hecium colou (hyaline o pale b own),
ascospo e and conidial sizes (Magnusson 1940, 1944; Ryan 2004; Kou e al. 2013;
Nimis 2016; Roux e al. 2016; Pauko e al. 2019; Kond a yuk e al. 2020; Zhang e
al. 2020, 2024a; Ash a e al. 2022; Zul iqa e al. 2022, 2023).
Seconda y me aboli es
Species o Lobo hallia can be ca ego ized in o wo g oups based on he
p esence o absence o seconda y me aboli es. Among species p oducing
seconda y me aboli es, no s ic ic acid is he p edominan and mos e-
quen ly encoun e ed compound (Ryan 2004; Roux 2012; Roux e al. 2016;
Kou e al. 2013; Pauko e al. 2019; Kond a yuk e al. 2020; Zhang e al.
2020; Ash a e al. 2022; Zul iqa e al. 2022, 2023). O he me aboli es in-
clude s ic ic acid (Kou e al. 2013; Moon and Kashiwadani 2009; Roux e
al. 2016; Pauko e al. 2019), cons ic ic acid (Ryan 2004; Kou e al. 2013),
conno s ic ic acid (Pauko e al. 2019), salazinic acid (Ryan 2004), and e -
penes (Galloway and Ledingham 2012; Roux e al. 2016).
All newly desc ibed species in his s udy con ain no s ic ic acid. Addi ion-
al seconda y me aboli es de ec ed include s ic ic acid in Lobo hallia pulch a,
L. ub a, and L. s ipi a a; c yp os ic ic acid ( eco ded o he i s ime in Lobo h-
allia) in L. benzilanensis, L. pulch a, and L. s ipi a a; gy opho ic acid exclusi ely
in L. ub a; and conno s ic ic acid, which equen ly occu ed as an accesso y
acid in all new species. Fu he mo e, hin-laye ch oma og aphy (TLC) analysis
o 11 known species (10 p e iously epo ed and one newly eco ded in China)
indica ed ha all species, excep L. c enula a, L. lobula a and L. subdi ac a,
con ain no s ic ic acid as he majo seconda y me aboli e (Kou e al. 2013;
Pauko e al. 2019; Zhang e al. 2024a). S ic ic acid was de ec ed in L. c as-
sima gina a; conno s ic ic acid was iden i ied in he species o L. alphoplaca,
L. c assima gina a, L. hadinii, L. p ae adiosa, L. p uinosa, and L. adiosa.
Phylogeny
Al hough he phylogene ic opology o Megaspo aceae om ou h ee-locus
da ase is la gely cong uen wi h he i e-locus esul s o Wheele e al. (2024),
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Lun Wang e al.: An o e iew o he genus Lobo hallia in China
and bo h place Lobo hallia as sis e o Teu oa, he s a is ical suppo o his e-
la ionship is conside ably lowe in ou analysis. Consequen ly, he ela ionship
be ween Lobo hallia and Teu oa emains unce ain and wa an s u he in es-
iga ion. Ou esul s con i m ha Lobo hallia comp ises h ee p ima y clades
(Clades I–III), consis en wi h p e ious s udies (Kou e al. 2013; Pauko e al.
2019; Zul iqa e al. 2022, 2023; Zhang e al. 2024a). Howe e , he e olu ion-
a y ela ionships among hese h ee clades emain un esol ed due o weak
suppo s a he backbone nodes (Fig. 1). The abo e esul s, o some ex en ,
suppo he assump ion o Wheele e al. (2024)—i Teu oa is accep ed a he
genus le el, hen a case could be made o e ec ing addi ional gene a wi hin
Lobo hallia. Whe he di e en clades o subclades o Lobo hallia wa an ec-
ogni ion as sepa a e gene a in he u u e will depend on comp ehensi e phylo-
gene ic and mo phological s udies wi hin he amily Megaspo aceae.
Fu he mo e, his s udy iden i ied six Lobo hallia species o which molec-
ula da a a e una ailable: L. chade audiana, L. gangwondoana, L. kumino ae,
L. lac eola, L. pla yca pa, and L. zog ii. Thei phylogene ic posi ions emain
un esol ed. Lobo hallia chade audiana is mo phologically mos simila o
L. ce noho skyana, di e ing p ima ily in he absence o seconda y me ab-
oli es (Clauzade and Vězda 1970; Roux 1977); i s phylogene ic posi ion is
en a i ely in e ed o belong o Clade III. Lobo hallia gangwondoana, cha -
ac e ized by a smoo h, sligh ly b ownish-g ay o cacao g ay hallus wi h in-
dis inc ma ginal lobes and dis an simple a eoles (Kond a yuk e al. 2020),
is en a i ely placed nea L. c assima gina a o L. benzilanensis wi hin Clade
I. Lobo hallia kumino ae, epo ed om Sibe ian Russia, is mo phologically
mo e o less iden ical o L. p ae adiosa based on i s o iginal desc ip ion
(Sedelniko a 1982). Fu he specimen examina ion is equi ed o de e mine
whe he hese wo axa a e conspeci ic. Lobo hallia lac eola, wi h i s ci cula
ou line, plica e ma gins, and lack o long and s aigh adial c acks (Pauko
e al. 2019), is en a i ely in e ed o belong o Clade III. Lobo hallia pla yca -
pa mo phologically esembles he subclade comp ising he h ee Pakis an
species, L. pul ina a, L. pakis anica and L. iqbalii. Lobo hallia zog ii, o igi-
nally desc ibed om Xinjiang P o ince, China, has a b own hallus, sho
ma ginal lobes, lecano ine apo hecia, and s ic ic acid (Pauko e al. 2019);
i s phylogene ic posi ion is en a i ely in e ed o be wi hin Clade I.
Acknowledgemen s
The au ho s hank D . Lulu Zhang om Shandong No mal Uni e si y o he
loan o specimens and pe mi ing DNA ex ac ion; D . Alexande G. Pauko ,
cu a o o U al Fede al Uni e si y, o p o iding he high-quali y mo phological
pho os o Lobo hallia zog ii.
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.
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Lun Wang e al.: An o e iew o he genus Lobo hallia in China
Use o AI
No use o AI was epo ed.
Funding
This s udy was unded by he Na ional Na u al Science Founda ion o China (32400005,
32570250), Yunnan Young & Eli e Talen s P ojec (YNWR-QNBJ-2020-224), Yunnan Fun-
damen al Resea ch P ojec s (202401AT070196), and he Second Tibe an Pla eau Scien-
i ic Expedi ion and Resea ch P og am (STEP) (2019QZKK0503).
Au ho con ibu ions
Concep ualiza ion: LSW, XYW, and YYZ; Da a cu a ion: LW and YYZ; Fo mal analysis: LW
and YSF; Funding acquisi ion: LSW, XYW, and YYZ; In es iga ion: LW and YSF; Me hodol-
ogy: LW and YSF; P ojec adminis a ion: LSW, XYW, and YYZ; Resou ces: LW and LSW;
So wa e: LW and YSF; Supe ision: XYW and YYZ; Valida ion: LW and YYZ; Visualiza-
ion: LW; W i ing – o iginal d a : LW and YYZ; Re iew, and edi ing: XYW and YYZ.
Au ho ORCIDs
Xin-Yu Wang h ps://o cid.o g/0000-0003-2166-6111
Yan-Yun Zhang h ps://o cid.o g/0000-0002-0902-5066
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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