Three new Diaporthe species from Shaanxi Province, China

Abstract

Diaporthe species (Sordariomycetes, Diaporthales) are often reported as important plant pathogens, saprobes and endophytes on a wide range of plant hosts. In this study, Diaporthe specimens were collected from symptomatic twigs and branches at the Huoditang Forest Farm in Shaanxi Province, China. Identification was done using a combination of morphology and comparison of DNA sequence data of the nuclear ribosomal internal transcribed spacer ( ITS), calmodulin (cal ), histone H3 ( his3 ), partial translation elongation factor-1α ( tef1 ) and β-tubulin ( tub2 ) gene regions. Three new Diaporthe species are proposed: D. albosinensis , D. coryli and D. shaanxiensis . All species are illustrated and their morphology and phylogenetic relationships with other Diaporthe species are discussed.

Full text

Taxonomy of Diaporthe 1
Three new Diaporthe species from
Shaanxi Province, China
Qin Yang1,2, Ning Jiang2, Cheng-Ming Tian2
1 Key Laboratory for Non-Wood Forest Cultivation and Conservation of the Ministry of Education, Central
South University of Forestry and Technology, Changsha 410004, China 2 e Key Laboratory for Silviculture
and Conservation of the Ministry of Education, Beijing Forestry University, Beijing 100083, China
Corresponding author: Cheng-Ming Tian (chengmt@bjfu.edu.cn)
Academic editor: D. Haelewaters|Received17 December 2019|Accepted 5 April 2020|Published 4 May2020
Citation: Yang Q, Jiang N, Tian C-M (2020) ree new Diaporthe species from Shaanxi Province, China. MycoKeys
67: 1–18. https://doi.org/10.3897/mycokeys.67.49483
Abstract
Diaporthe species (Sordariomycetes, Diaporthales) are often reported as important plant pathogens, sap-
robes and endophytes on a wide range of plant hosts. In this study, Diaporthe specimens were collected
from symptomatic twigs and branches at the Huoditang Forest Farm in Shaanxi Province, China. Identi-
cation was done using a combination of morphology and comparison of DNA sequence data of the nu-
clear ribosomal internal transcribed spacer (ITS), calmodulin (cal), histone H3 (his3), partial translation
elongation factor-1α (tef1) and β-tubulin (tub2) gene regions. ree new Diaporthe species are proposed:
D. albosinensis, D. coryli and D. shaanxiensis. All species are illustrated and their morphology and phylo-
genetic relationships with other Diaporthe species are discussed.
Keywords
Diaporthaceae, Dieback, DNA phylogeny, Systematics, Taxonomy
Introduction
Diaporthe species (Sordariomycetes, Diaporthales) are associated with a wide range of
plant hosts as pathogens, endophytes or saprobes of crops, ornamentals and forest trees
(Murali et al. 2006, Rossman et al. 2007, Garcia-Reyne et al. 2011, Gomes et al. 2013,
Udayanga et al. 2015, Dissanayake et al. 2017, Guarnaccia and Crous2017,2018,
Copyright Qin Yang et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
MycoKeys 67: 1–18 (2020)
doi: 10.3897/mycokeys.67.49483
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Qin Yang et al. / MycoKeys 67: 1–18 (2020)
2
Wijayawardene et al. 2017, Yang et al. 2017a, b, 2018, Fan et al. 2018, Guarnaccia et
al. 2018). e sexual morph of Diaporthe is characterised by immersed ascomata and
an erumpent pseudostroma with elongated perithecial necks. Asci are unitunicate,
clavate to cylindrical. Ascospores are fusoid, ellipsoid to cylindrical, hyaline, biseriate
to unise riate in the ascus, sometimes with appendages (Udayanga et al. 2011). e
asexual morph is characterised by ostiolate conidiomata, with cylindrical phialides
producing three types of hyaline, aseptate conidia (Udayanga et al. 2011, Gomes
etal.2013).
Species identication in Diaporthe has traditionally been based on host associa-
tion, morphology and culture characteristics (Mostert et al. 2001, Santos and Phillips
2009, Udayanga et al. 2011), resulting in the description of over 200 species (Hyde et
al. 2020). Multiple species of Diaporthe can colonise a single host and one species can
be associated with dierent hosts (Santos and Phillips 2009, Diogo et al. 2010, Santos
et al. 2011, Gomes et al. 2013). In addition, considerable within-species variability
of phenotypic characters has been reported (Rehner and Uecker 1994, Mostert et al.
2001, Udayanga et al. 2011). us, a polyphasic taxonomic approach, based on multi-
locus DNA data, morphology and ecology, has been increasingly employed for species
boundaries in the genus Diaporthe (Gomes et al. 2013, Huang et al. 2013, 2015,
Udayanga et al. 2014a, b, 2015, Fan et al. 2015, Du et al. 2016, Gao et al. 2016, 2017,
Guarnaccia and Crous 2017, Guarnaccia et al. 2018, Long et al. 2019).
Huoditang is located in the middle part of the southern slope of the Qinling
Mountains at 33°18'~33°28'N, 108°21'~108°29'E. It belongs to the transitional zone
of the northern subtropical and warm temperate zone in China. e terrain is com-
plex and the climate is changeable (Zhang and Cao 2007). e plant communities are
complex and, as a result, species diversity of fungi in the forest area is high (Zhang and
Cao 2007). During trips to collect forest pathogens causing dieback in Shaanxi Prov-
ince, cankered branches with typical Diaporthe fruiting bodies were investigated and
sampled. e aim of the present study was to identify these fungi, based on modern
polyphasic taxonomic concepts.
Materials and methods
Isolates
Fresh specimens of Diaporthe were collected from symptomatic twigs or branches in
Shaanxi Province (Table 1). Isolates were ob tained by removing a mucoid spore mass
from conidiomata and spreading the suspen sion on the surface of 1.8% potato dex-
trose agar (PDA) in a 9 cm diam. Petri dish. Petri dishes were incubated at 25 °C until
spores germinated. Single germinating conidia were transferred on to new PDA plates,
which were kept at 25 °C in the dark. Specimens are deposited in the Museum of the
Beijing Forestry University (BJFC). Axenic cultures are maintained in the China For-
estry Culture Collection Centre (CFCC).

Taxonomy of Diaporthe 3
Table 1. Isolates and GenBank accession numbers used in the phylogenetic analyses of Diaporthe.
Species Isolate Host Location GenBank accession numbers
ITS cal his3 tef1 tub2
D. acericola MFLUCC
17-0956
Acer negundo Italy KY964224 KY964137 NA KY964180 KY964074
D. acerigena CFCC 52554 Acer tataricum China MH121489 MH121413 MH121449 MH121531 NA
D. albosinensis CFCC 53066 Betula
albosinensis
China MK432659 MK442979 MK443004 MK578133 MK578059
CFCC 53067 Betula
albosinensis
China MK432660 MK442980 MK443005 MK578134 MK578060
D. alnea CBS 146.46 Alnus sp. Netherlands KC343008 KC343250 KC343492 KC343734 KC343976
D. ambigua CBS 114015 Pyrus communis South Africa KC343010 KC343252 KC343494 KC343736 KC343978
D. anacardii CBS 720.97 Anacardium
occidentale
East Africa KC343024 KC343266 KC343508 KC343750 KC343992
D. angelicae CBS 111592 Heracleum
sphondylium
Austria KC343027 KC343269 KC343511 KC343753 KC343995
D. apiculatum CGMCC
3.17533
Camellia sinensis China KP267896 NA NA KP267970 KP293476
D. aquatica IFRDCC 3051 Aquatic habitat China JQ797437 NA NA NA NA
D. arctii CBS 139280 Arctium lappa Austria KJ590736 KJ612133 KJ659218 KJ590776 KJ610891
D. aseana MFLUCC
12-0299a
Unknown dead
leaf
Thailand KT459414 KT459464 NA KT459448 KT459432
D. asheicola CBS 136967 Vaccinium ashei Chile KJ160562 KJ160542 NA KJ160594 KJ160518
D. baccae CBS 136972 Vaccinium
corymbosum
Italy KJ160565 NA MF418264 KJ160597 NA
D. beilharziae BRIP 54792 Indigofera
australis
Australia JX862529 NA NA JX862535 KF170921
D. benedicti BPI 893190 Salix sp. USA KM669929 KM669862 NA KM669785 NA
D. betulae CFCC 50469 Betula
platyphylla
China KT732950 KT732997 KT732999 KT733016 KT733020
D. betulina CFCC 52560 Betula albo-
sinensis
China MH121495 MH121419 MH121455 MH121537 MH121577
D. bicincta CBS 121004 Juglans sp. USA KC343134 KC343376 KC343618 KC343860 KC344102
D. caryae CFCC 52563 Carya illinoensis China MH121498 MH121422 MH121458 MH121540 MH121580
D. cassines CPC 21916 Cassine peragua South Africa KF777155 NA NA KF777244 NA
D. celeris CPC 28262 Vitis vinifera Czech
Republic
MG281017 MG281712 MG281363 MG281538 MG281190
D. cercidis CFCC 52565 Cercis chinensis China MH121500 MH121424 MH121460 MH121542 MH121582
D. chamaeropis CBS 454.81 Chamaerops
humilis
Greece KC343048 KC343290 KC343532 KC343774 KC344016
D. charlesworthii BRIP 54884m Rapistrum
rugostrum
Australia KJ197288 NA NA KJ197250 KJ197268
D. chensiensis CFCC 52567 Abies chensiensis China MH121502 MH121426 MH121462 MH121544 MH121584
D. cichorii MFLUCC
17-1023
Cichorium
intybus
Italy KY964220 KY964133 NA KY964176 KY964104
D. cinnamomi CFCC 52569 Cinnamomum
sp.
China MH121504 NA MH121464 MH121546 MH121586
D. citriasiana CGMCC
3.15224
Citrus unshiu China JQ954645 KC357491 KJ490515 JQ954663 KC357459
D. citrichinensis CGMCC
3.15225
Citrus sp. China JQ954648 KC357494 NA JQ954666 NA
D. compactum CGMCC
3.17536
Camellia sinensis China KP267854 NA KP293508 KP267928 KP293434
D. conica CFCC 52571 Alangium
chinense
China MH121506 MH121428 MH121466 MH121548 MH121588
D. coryli CFCC 53083 Corylus
mandshurica
China MK432661 MK442981 MK443006 MK578135 MK578061
CFCC 53084 Corylus
mandshurica
China MK432662 MK442982 MK443007 MK578136 MK578062
D. cucurbitae CBS 136.25 Arctium sp. Unknown KC343031 KC343273 KC343515 KC343757 KC343999
D. cuppatea CBS 117499 Aspalathus
linearis
South Africa KC343057 KC343299 KC343541 KC343783 KC344025

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
4
Species Isolate Host Location GenBank accession numbers
ITS cal his3 tef1 tub2
D. cynaroidis CBS 122676 Protea cynaroides South Africa KC343058 KC343300 KC343542 KC343784 KC344026
D. cytosporella FAU461 Citrus limon Italy KC843307 KC843141 NA KC843116 KC843221
D. discoidispora ZJUD89 Citrus unshiu China KJ490624 NA KJ490566 KJ490503 KJ490445
D. dorycnii MFLUCC
17-1015
Dorycnium
hirsutum
Italy KY964215 NA NA KY964171 KY964099
D. elaeagni-glabrae CGMCC
3.18287
Elaeagnus glabra China KX986779 KX999281 KX999251 KX999171 KX999212
D. endophytica CBS 133811 Schinus
terebinthifolius
Brazil KC343065 KC343307 KC343549 KC343791 KC343065
D. eres AR5193 Ulmus sp. Germany KJ210529 KJ434999 KJ420850 KJ210550 KJ420799
D. eucalyptorum CBS 132525 Eucalyptus sp. Australia NR120157 NA NA NA NA
D. foeniculacea CBS 123208 Foeniculum
vulgare
Portugal KC343104 KC343346 KC343588 KC343830 KC344072
D. fraxini-
angustifoliae
BRIP 54781 Fraxinus
angustifolia
Australia JX862528 NA NA JX862534 KF170920
D. fraxinicola CFCC 52582 Fraxinus
chinensis
China MH121517 MH121435 NA MH121559 NA
D. fructicola MAFF 246408 Passiflora edulis ×
P. edulis f.
flavicarpa
Japan LC342734 LC342738 LC342737 LC342735 LC342736
D. fusicola CGMCC
3.17087
Lithocarpus
glabra
China KF576281 KF576233 NA KF576256 KF576305
D. garethjonesii MFLUCC
12-0542a
Unknown dead
leaf
Thailand KT459423 KT459470 NA KT459457 KT459441
D. guangxiensis JZB320094 Vitis vinifera China MK335772 MK736727 NA MK523566 MK500168
D. helicis AR5211 Hedera helix France KJ210538 KJ435043 KJ420875 KJ210559 KJ420828
D. heterophyllae CBS 143769 Acacia
heterohpylla
France MG600222 MG600218 MG600220 MG600224 MG600226
D. hubeiensis JZB320123 Vitis vinifera China MK335809 MK500235 NA MK523570 MK500148
D. incompleta CGMCC
3.18288
Camellia sinensis China KX986794 KX999289 KX999265 KX999186 KX999226
D. inconspicua CBS 133813 Maytenus
ilicifolia
Brazil KC343123 KC343365 KC343607 KC343849 KC344091
D. infecunda CBS 133812 Schinus
terebinthifolius
Brazil KC343126 KC343368 KC343610 KC343852 KC344094
D. juglandicola CFCC 51134 Juglans
mandshurica
China KU985101 KX024616 KX024622 KX024628 KX024634
D. kadsurae CFCC 52586 Kadsura
longipedunculata
China MH121521 MH121439 MH121479 MH121563 MH121600
D. litchicola BRIP 54900 Litchi chinensis Australia JX862533 NA NA JX862539 KF170925
D. lusitanicae CBS 123212 Foeniculum
vulgare
Portugal KC343136 KC343378 KC343620 KC343862 KC344104
D. masirevicii BRIP 57892a Helianthus
annuus
Australia KJ197277 NA NA KJ197239 KJ197257
D. middletonii BRIP 54884e Rapistrum
rugostrum
Australia KJ197286 NA NA KJ197248 KJ197266
D. millettiae GUCC9167 Millettia
reticulata
China MK398674 MK502086 NA MK480609 MK502089
D. miriciae BRIP 54736j Helianthus
annuus
Australia KJ197282 NA NA KJ197244 KJ197262
D. musigena CBS 129519 Musa sp. Australia KC343143 KC343385 KC343627 KC343869 KC344111
D. neilliae CBS 144.27 Spiraea sp. USA KC343144 KC343386 KC343628 KC343870 KC344112
D. neoarctii CBS 109490 Ambrosia trifida USA KC343145 KC343387 KC343629 KC343871 KC344113
D. nothofagi BRIP 54801 Nothofagus
cunninghamii
Australia JX862530 NA NA JX862536 KF170922
D. novem CBS 127270 Glycine max Croatia KC343155 KC343397 KC343640 KC343881 KC344123
D. oraccinii CGMCC
3.17531
Camellia sinensis China KP267863 NA KP293517 KP267937 KP293443
D. ovalispora ICMP20659 Citrus limon China KJ490628 NA KJ490570 KJ490507 KJ490449
D. ovoicicola CGMCC
3.17093
Citrus sp. China KF576265 KF576223 NA KF576240 KF576289

Taxonomy of Diaporthe 5
Species Isolate Host Location GenBank accession numbers
ITS cal his3 tef1 tub2
D. osmanthi GUCC9165 Osmanthus
fragrans
China MK398675 MK502087 NA MK480610 MK502090
D. padina CFCC 52590 Padus racemosa China MH121525 MH121443 MH121483 MH121567 MH121604
D. pandanicola MFLU
18-0006
Pandanus sp. Thailand MG646974 NA NA NA MG646930
D. pascoei BRIP 54847 Persea americana Australia JX862532 NA NA JX862538 KF170924
D. passifloricola CBS 141329 Passiflora foetida Malaysia KX228292 NA KX228367 NA KX228387
D. perseae CBS 151.73 Persea gratissima Netherlands KC343173 KC343415 KC343657 KC343899 KC344141
D. pescicola MFLUCC
16-0105
Prunus persica China KU557555 KU557603 NA KU557623 KU557579
D. phaseolorum AR4203 Phaseolus
vulgaris
USA KJ590738 NA KJ659220 NA KP004507
D. podocarpi-
macrophylli
CGMCC
3.18281
Podocarpus
macrophyllus
China KX986774 KX999278 KX999246 KX999167 KX999207
D.
pseudomangiferae
CBS 101339 Mangifera indica Dominican
Republic
KC343181 KC343423 KC343665 KC343907 KC344149
D.
pseudophoenicicola
CBS 462.69 Phoenix
dactylifera
Spain KC343184 KC343426 KC343668 KC343910 KC344152
D. psoraleae-
pinnatae
CBS 136413 Psoralea pinnata South Africa KF777159 NA NA NA KF777252
D. pulla CBS 338.89 Hedera helix Yugoslavia KC343152 KC343394 KC343636 KC343878 KC344120
D. racemosae CBS 143770 Euclea racemosa South Africa MG600223 MG600219 MG600221 MG600225 MG600227
D. ravennica MFLUCC
15-0479
Tamarix sp. Italy KU900335 NA NA KX365197 KX432254
D. rhusicola CBS 129528 Rhus pendulina South Africa JF951146 KC843124 NA KC843100 KC843205
D. rosae MFLU
17-1550
Rosa sp. Thailand MG828894 NA NA NA MG843878
D. rosicola MFLU
17-0646
Rosa sp. UK MG828895 NA NA MG829270 MG843877
D. rudis AR3422 Laburnum
anagyroides
Austria KC843331 KC843146 NA KC843090 KC843177
D. sackstonii BRIP 54669b Helianthus
annuus
Australia KJ197287 NA NA KJ197249 KJ197267
D. salicicola BRIP 54825 Salix purpurea Australia JX862531 NA NA JX862537 JX862531
D. sambucusii CFCC 51986 Sambucus
williamsii
China KY852495 KY852499 KY852503 KY852507 KY852511
D. schini CBS 133181 Schinus
terebinthifolius
Brazil KC343191 KC343433 KC343675 KC343917 KC344159
D. schoeni MFLU
15-1279
Schoenus
nigricans
Italy KY964226 KY964139 NA KY964182 KY964109
D. sennicola CFCC 51634 Senna
bicapsularis
China KY203722 KY228873 KY228879 KY228883 KY228889
D. serafiniae BRIP 55665a Helianthus
annuus
Australia KJ197274 NA NA KJ197236 KJ197254
D. shaanxiensis CFCC 53106 on branches of
liana
China MK432654 MK442976 MK443001 MK578130 NA
CFCC 53107 on branches of
liana
China MK432655 MK442977 MK443002 MK578131 NA
D. siamensis MFLUCC
10-573a
Dasymaschalon
sp.
Thailand JQ619879 NA NA JX275393 JX275429
D. sojae FAU635 Glycine max USA KJ590719 KJ612116 KJ659208 KJ590762 KJ610875
D. sterilis CBS 136969 Vaccinium
corymbosum
Italy KJ160579 KJ160548 MF418350 KJ160611 KJ160528
D. stictica CBS 370.54 Buxus
sampervirens
Italy KC343212 KC343454 KC343696 KC343938 KC344180
D. subclavata ICMP20663 Citrus unshiu China KJ490587 NA KJ490529 KJ490466 KJ490408
D. subcylindrospora MFLU
17-1195
Salix sp. China MG746629 NA NA MG746630 MG746631
D. subellipicola MFLU
17-1197
on dead wood China MG746632 NA NA MG746633 MG746634

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
6
Species Isolate Host Location GenBank accession numbers
ITS cal his3 tef1 tub2
D. subordinaria CBS 464.90 Plantago
lanceolata
New
Zealand
KC343214 KC343456 KC343698 KC343940 KC344182
D.
tectonendophytica
MFLUCC
13-0471
Tectona grandis China KU712439 KU749354 NA KU749367 KU749354
D. tectonigena MFLUCC
12-0767
Tectona grandis China KU712429 KU749358 NA KU749371 KU743976
D. terebinthifolii CBS 133180 Schinus
terebinthifolius
Brazil KC343216 KC343458 KC343700 KC343942 KC344184
D. ternstroemia CGMCC
3.15183
Ternstroemia
gymnanthera
China KC153098 NA NA KC153089 NA
D. thunbergii MFLUCC
10-576a
Thunbergia
laurifolia
Thailand JQ619893 JX197440 NA JX275409 JX275449
D. tibetensis CFCC 51999 Juglandis regia China MF279843 MF279888 MF279828 MF279858 MF279873
D. ueckerae FAU656 Cucumis melo USA KJ590726 KJ612122 KJ659215 KJ590747 KJ610881
D. ukurunduensis CFCC 52592 Acer
ukurunduense
China MH121527 MH121445 MH121485 MH121569 NA
D. unshiuensis CFCC 52594 Carya illinoensis China MH121529 MH121447 MH121487 MH121571 MH121606
D. vaccinii CBS 160.32 Oxycoccus
macrocarpos
USA KC343228 KC343470 KC343712 KC343954 KC344196
D. velutina CGMCC
3.18286
Neolitsea sp. China KX986790 NA KX999261 KX999182 KX999223
D. viniferae JZB320071 Vitis vinifera China MK341551 MK500107 NA MK500119 MK500112
D. xishuangbanica CGMCC
3.18282
Camellia sinensis China KX986783 NA KX999255 KX999175 KX999216
D. yunnanensis CGMCC
3.18289
Coffea sp. China KX986796 KX999290 KX999267 KX999188 KX999228
Diaporthella
corylina
CBS 121124 Corylus sp. China KC343004 KC343246 KC343488 KC343730 KC343972
Newly sequenced material is indicated in bold type. NA, not applicable.
Morphological analysis
Morphological observations of the asexual morph in the natural environment were
based on features of the fruiting bodies produced on infected plant tissues and micro-
morphology, supplemented by cultural characteristics. Conidiomata from tree barks
were sectioned by hand, using a double-edged blade and structures were observed un-
der a dissecting microscope. e gross morphology of fruiting bodies was recorded
using a Leica stereomicroscope (M205 FA). Fungal structures were mounted in clear
lactic acid and micromorphological characteristics were examined at 1000× magnica-
tion using a Leica compound microscope (DM 2500) with dierential interference
contrast (DIC) optics. irty measurements of each structure were determined for
each collection. Colony characters and pigment production on PDA were noted after
10 d. Colony colours were described according to Rayner (1970).
DNA extraction, PCR amplification and sequencing
Genomic DNA was extracted from colonies grown on cellophane-covered PDA, us-
ing the CTAB [cetyltrimethylammonium bromide] method (Doyle and Doyle 1990).
PCR amplications of phylogenetic markers were done using the same primer pairs

Taxonomy of Diaporthe 7
and conditions as in Yang et al. (2018). PCR products were assayed via electrophoresis
in 2% agarose gels. DNA sequencing was performed using an ABI PRISM 3730XL
DNA Analyzer with a BigDye Terminater Kit v.3.1 (Invitrogen, USA) at the Shanghai
Invitrogen Biological Technology Company Limited (Beijing, China).
Phylogenetic analyses
e quality of our amplied nucleotide sequences was checked and combined by Seq-
Man v.7.1.0 and reference sequences were retrieved from the National Center for Bio-
technology Information (NCBI), based on recent publications on the genus Diaporthe
(Guarnaccia et al. 2018, Yang et al. 2018, Long et al. 2019). Sequences were aligned
using MAFFT v. 7.310 (http://mat.cbrc.jp/alignment/server/index.html) (Katoh and
Standley 2016) and manually corrected using Bioedit 7.0.9.0 (Hall 1999). e best-t
nucleotide substitution models for each gene were selected using jModelTest v. 2.1.7
(Darriba et al. 2012) under the Akaike Information Criterion.
Phylogenetic analyses of the combined gene regions were performed using Maxi-
mum-Likelihood (ML) and Bayesian Inference (BI) methods. ML was conducted us-
ing PhyML v. 3.0 (Guindon et al. 2010), with 1000 bootstrap replicates. BI was per-
formed using a Markov Chain Monte Carlo (MCMC) algorithm in MrBayes v. 3.0b4
(Ronquist and Huelsenbeck 2003). Two MCMC chains, started from random trees for
1,000,000 generations and trees, were sampled every 100th generation, resulting in a
total of 10,000 trees. e rst 25% of trees were discarded as burn-in of each analysis.
Branches with signicant Bayesian Posterior Probabilities (BPP) were estimated in the
remaining 7500 trees. Phylogenetic trees were viewed with FigTree v.1.3.1 (Rambaut
and Drummond 2010) and processed by Adobe Illustrator CS5. Alignment and trees
were deposited in TreeBASE (submission ID: S25522). e nucleotide sequence data
of the new taxa have been deposited in GenBank (Table 1).
Results
Phylogenetic analyses
e ve-gene sequence dataset (ITS, cal, his3, tef1 and tub2) was analysed to infer the
interspecic relationships within Diaporthe. e dataset consisted of 124 sequences
including the outgroup, Diaporthella corylina (culture CBS 121124). A total of 2555
characters including gaps (505 for ITS, 513 for cal, 528 for his3, 475 for tef1 and 522
for tub2) were included in the phylogenetic analysis. e best nucleotide substitution
model for ITS, his3 and tub2 was TrN+I+G, while HKY+I+G was selected for both cal
and tef1. e topologies resulting from ML and BI analyses of the concatenated dataset
were congruent (Fig. 1). Isolates from Shaanxi Province formed three individual clades
representing three undescribed species.

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
8
Figure 1. Phylogram of Diaporthe resulting from a maximum likelihood analysis based on combined
ITS, cal, his3, tef1 and tub2. Numbers above the branches indicate ML bootstraps (left, ML BS ≥ 50%)
and Bayesian Posterior Probabilities (right, BPP ≥ 0.90). e tree is rooted with Diaporthella corylina.
Isolates in current study are in blue. “-” indicates ML BS < 50% or BI PP < 0.90.

Taxonomy of Diaporthe 9
Taxonomy
Diaporthe albosinensis C.M. Tian & Q. Yang, sp. nov.
MycoBank No: 829518
Fig. 2
Diagnosis. Distinguished from D. fraxinicola in having shorter conidiophores and
longer beta conidia.
Etymology. Named after the host plant, Betula albosinensis, from which the holo-
type was collected.
Description. Conidiomata pycnidial, conical, immersed in bark, solitary to ag-
gregated, erumpent through the bark surface, with a solitary undivided locule.
Ectostromatic disc yellowish to brown, one ostiole per disc. Ostiole medium black,
Figure 1. Continued.

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
10
up to the level of disc. Locule undivided, (280–)290–375(–380) µm diam. Conidi-
ophores (6–)8.5–13(–14.5) × (1.5–)2–2.5 µm, hyaline, cylindrical, smooth, phialidic,
unbranched, straight or slightly curved. Alpha conidia hyaline, aseptate, fusiform,
0–1-guttulate, (7–)8–10(–11) × 2.5–3 µm. Beta conidia hyaline, aseptate, liform,
Figure 2. Diaporthe albosinensis on Betula albosinensis (BJFC-S1670). A Habit of conidiomata in wood
B transverse section of conidiomata C longitudinal section through conidiomata D conidiogenous cells
attached with beta conidia E conidiogenous cells attached with alpha conidia F beta conidia. Scale bars:
200 µm (B–C); 20 µm (D, F ); 10 µm (E).

Taxonomy of Diaporthe 11
straight or slightly curved, eguttulate, base subtruncate, tapering towards one apex,
(24–)25.5–30(–32)× 1–1.5 µm.
Culture characters. Cultures incubated on PDA at 25 °C in the dark. Colony
originally at with white felted aerial mycelium, becoming light brown due to pig-
ment formation, conidiomata irregularly distributed over agar surface, with yellowish
conidial drops exuding from the ostioles.
Specimens examined. C. Shaanxi Province: Ningshan County, Huoditang
Forest Farm, 33°28'25"N, 108°29'39"E, on branches of Betula albosinensis, 10 July
2018, N. Jiang (holotype BJFC-S1670; ex-type living culture: CFCC 53066; living
culture: CFCC 53067).
Notes. Two isolates, representing D. albosinensis, are retrieved in a well-supported
clade (ML BS/BPP=100/1) and appear most closely related to D. fraxinicola (Fig. 1).
Diaporthe albosinensis can be distinguished from D. fraxinicola, based on tef1 and tub2
loci (3/335 in tef1 and 19/429 in tub2). Morphologically, D. albosinensis diers from
D. fraxinicola in having shorter conidiophores (8.5–13 vs. 10.5–17.5 µm) and longer
beta conidia (25.5–30 vs. 19–29.5 µm) (Yang et al. 2018).
Diaporthe coryli C.M. Tian & Q. Yang, sp. nov.
MycoBank No: 829520
Fig. 3
Diagnosis. Distinguished from D. ukurunduensis and D. citrichinensis in having larger
alpha conidia.
Etymology. Named after the genus of the host plant from which the holotype was
collected, Corylus.
Description. Conidiomata pycnidial, conical to spherical, immersed in the host
bark, erumpent from surface of host branches, scattered, 950–1200 × 420–650 µm
diam., covered by orange discharged conidial masses at maturity, usually conspicuous.
Ectostromatic disc inconspicuous. Central column beneath the disc more or less conical,
bright yellow. Conidiophores reduced to conidiogenous cells. Conidiogenous cells cylin-
drical, hyaline, smooth, unbranched, tapering towards the apex, (8.5–)10–12(–13) ×
(2–)2.5–3 µm. Alpha conidia hyaline, aseptate, fusiform, multiguttulate, rarely 2-gut-
tulate, (10.5–)11.5–13(–13.5) × 3–3.5 µm. Beta conidia not observed.
Culture characters. Cultures incubated on PDA at 25 °C in the dark. Colony at,
felty with thick texture at the marginal area, with thin texture in the centre, producing
beige pigment after 7–10 d. Aerial mycelium white, dense, conidiomata distributed in
the centre, with translucent conidial drops exuding from the ostioles.
Specimens examined. CHINA. Shaanxi Province: Ningshan County, Huodi-
tang Forest Farm, 33°28'26"N, 108°29'40"E, on branches of Corylus mandshurica,
10 July 2018, N. Jiang (holotype BJFC-S1671; ex-type living culture: CFCC 53083);
33°28'26"N, 108°29'38"E, on branches of Corylus mandshurica, 10 July 2018, N.
Jiang (paratype BJFC-S1672; living culture: CFCC 53084).

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
12
Notes. We generated sequences for two isolates of D. coryli, CFCC 53083 and
CFCC 53084. is new species is phylogenetically most closely related to D. uku-
runduensis and D. citrichinensis (Fig. 1). Diaporthe coryli can be distinguished from
D. ukurunduensis, based on ITS, his3 and tef1 loci (8/467 in ITS, 1/460 in his3 and
1/336 in tef1); and from D. citrichinensis based on tef1 and tub2 loci (4/335 in tef1
and 25/428 in tub2). Morphologically, D. coryli can be distinguished from both D.
ukurunduensis (11.5–13 × 3–3.5 vs. 5–6 × 2–3 µm) and D. citrichinensis (11.5–13×
3–3.5 vs. 5.5–9 × 1.5–2.5 µm) in having larger alpha conidia (Huang et al. 2013,
Gao et al. 2016).
Figure 3. Diaporthe coryli on Corylus mandshurica (BJFC-S1671). A , B Habit of conidiomata in wood
C transverse section of conidiomata D longitudinal section through conidiomata E conidiogenous cells
attached with alpha conidia F alpha conidia. Scale bars: 500 µm (B–D); 10 µm (E); 20 µm (F).

Taxonomy of Diaporthe 13
Diaporthe shaanxiensis C.M. Tian & Q. Yang, sp. nov.
MycoBank No: 829527
Fig. 4
Diagnosis. Distinguished from D. aquatica and D. incompleta in having longer
beta conidia.
Etymology. Named after Province Shaanxi, where the holotype was collected.
Description. Conidiomata pycnidial, immersed in bark, scattered, erumpent
through the bark surface, discoid, with a solitary undivided locule. Ectostromatic disc
yellowish to pale brown, one ostiole per disc, usually conspicuous, (485–)500–687(–
Figure 4. Diaporthe shaanxiensis on liana (BJFC-S1674). A, B Habit of conidiomata on twig C trans-
verse section through conidiomata D longitudinal section through conidiomata E conidiogenous cells
attached with beta conidia F beta conidia. Scale bars: 200 µm (B–D); 10 µm (E, F).

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
14
695) µm diam. Locule circular, undivided, (500–)526–765(–792) µm diam. Conidi-
ophores reduced to conidiogenous cells. Conidiogenous cells hyaline, cylindrical, un-
branched, slightly curved, tapering towards the apex, (12.5–)14.5–17(–18) × 1–1.5(–
2) µm. Alpha conidia not observed. Beta conidia hyaline, aseptate, liform, straight to
curved, eguttulate, (35.5–)37–47.5(–50) × 1 µm.
Culture characters. Cultures incubated on PDA at 25 °C in the dark. Colony
originally at with white uy aerial mycelium, becoming pale brown with pigment,
with visible solitary conidiomata at maturity.
Specimens examined. CHINA. Shaanxi Province: Ningshan County, Huodi-
tang Forest Farm, 33°28'25"N, 108°29'39"E, on branch of liana, 10 July 2018, N.
Jiang (holotype BJFC-S1674; ex-type living culture: CFCC 53106); 33°28'24"N,
108°29'38"E, on branch of liana, 10 July 2018, N. Jiang (Paratype BJFC-S1675; liv-
ing culture: CFCC 53107).
Notes. In the combined tree, D. shaanxiensis is a distinct clade with maximum
support and it appears to be most closely related to D. aquatica and D. incompleta (Fig.
1). Diaporthe shaanxiensis can be distinguished from D. aquatica by a 17 nt dierence
in the ITS region. For D. aquatica, only ITS sequences are available in NCBI GenBank
(Hu et al. 2012). e new species can be distinguished from D. incompleta, based
on ITS, cal, his3 and tef1 (24/454 in ITS, 14/443 in cal, 66/468 in his3 and 24/311
in tef1). Morphologically, D. shaanxiensis diers from both D. aquatica (37–47.5 vs.
9–12.5 µm) and D. incompleta (37–47.5 vs. 19–44 µm) in having longer beta conidia
(Gao et al. 2016, 2017).
Discussion
In this study, an investigation of forest pathogens from Huoditang in Shaanxi Province
was carried out and Diaporthe canker was observed as a common disease. Identica-
tion of our collections was conducted, based on isolates from fruiting bodies using ve
combined loci (ITS, cal, his3, tef1 and tub2), as well as morphological characters. ree
new Diaporthe species were described. ese are D. albosinensis sp. nov., D. coryli sp.
nov. and D. shaanxiensis sp. nov.
Diaporthe albosinensis is associated with Betula albosinensis. us far, six Diaporthe
species have been reported from Betula. ese are D. alleghaniensis, D. betulae, D. betu-
licola, D. betulina, D. eres and D. melanocarpa (Kobayashi 1970, Gomes et al. 2013, Du
et al. 2016, Yang et al. 2018). Morphologically, D. albosinensis diers from D. betulae
(600–1250 µm), D. betulicola (700–1300 µm) and D. betulina (670–905 µm) in hav-
ing smaller locules (Du et al. 2016, Yang et al. 2018); and from D. alleghaniensis (5–8×
1.5–2 µm) and D. eres (6.5–8.5 × 3–4 µm) in having larger alpha conidia (Arnold 1967,
Anagnostakis 2007, Gomes et al. 2013). In addition, our phylogenetic reconstruction
of a ve-locus dataset adds support for the new species, although no sequence data
are currently available for D. alleghaniensis, D. betulicola and D. melanocarpa (Fig. 1).
Interestingly, D. melanocarpa is found on dierent plant hosts; it was described from

Taxonomy of Diaporthe 15
Pyrus melanocarpa in London and then recorded from Amelanchier, Betula and Cornus
(Dearness 1926, Wehmeyer 1933, Kobayashi 1970). Diaporthe coryli is characterised by
the ostiole with orange discharged conidial masses and a yellow central column (Fig. 3).
Diaporthe shaanxiensis was found on branches of liana with an obvious ostiole per disc
and characterised by hyaline, liform beta conidia. Alpha conidia were found neither in
the natural environment nor in culture for this species.
Species delimitation of Diaporthe has improved considerably by using a combina-
tion of morphological, cultural, phytopathological and molecular phylogenetic analy-
ses (Udayanga et al. 2014a, b, 2015, Fan et al. 2015, Gao et al. 2017, Guarnaccia and
Crous 2017, Hyde et al. 2017, 2020, Guarnaccia et al. 2018, Yang et al. 2018, Long
et al. 2019). As a result, many Diaporthe canker diseases and new species have been
discovered and reported from all over the world and also in China. e descriptions
and molecular data of Diaporthe species represent an important resource for plant pa-
thologists, plant quarantine ocials and taxonomists.
Acknowledgements
is study is nanced by the Research Foundation of Education Bureau of Hunan
Province, China (Project No.: 19B608) and the introduction of talent research start-
up fund project of CSUFT (Project No.: 2019YJ025). We are grateful to Chungen
Piao, Minwei Guo (China Forestry Culture Collection Center, Chinese Academy of
Forestry, Beijing) and reviewers Lu Quan and Jadson Bezerra.
References
Crous PW, Gams W, Stalpers JA, Robert V, Stegehuis G (2004) MycoBank: an online initiative
to launch mycology into the 21st century. Studies in Mycology 50: 19–22.
Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more models, new heuris-
tics and parallel computing. Nature Methods 9: 772. https://doi.org/10.1038/nmeth.2109
Dearness J (1926) New and noteworthy fungi. Mycologia 18: 236–255. https://doi.org/10.10
80/00275514.1926.12020515
Diogo E, Santos JM, Phillips AJ (2010) Phylogeny, morphology and pathogenicity of Diaporthe
and Phomopsis species on almond in Portugal. Fungal Diversity 44: 107–115. https://doi.
org/10.1007/s13225-010-0057-x
Dissanayake AJ, Phillips AJL, Hyde KD, Yan JY, Li XH (2017) e current status of species in
Diaporthe. Mycosphere 8: 1106–1156. https://doi.org/10.5943/mycosphere/8/5/5
Doyle JJ, Doyle JL (1990) Isolation of plant DNA from fresh tissue. Focus 12: 13–15. https://
doi.org/10.2307/2419362
Du Z, Fan XL, Hyde KD, Yang Q, Liang YM, Tian CM (2016) Phylogeny and morphology
reveal two new species of Diaporthe from Betula spp. in China. Phytotaxa 269: 90–102.
https://doi.org/10.11646/phytotaxa.269.2.2

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
16
Fan XL, Hyde KD, Udayanga D, Wu XY, Tian CM (2015) Diaporthe rostrata, a novel ascomy-
cete from Juglans mandshurica associated with walnut dieback. Mycological Progress 14:
1–8. https://doi.org/10.1007/s11557-015-1104-5
Fan XL, Yang Q, Bezerra JDP, Alvarez LV, Tian CM (2018) Diaporthe from walnut tree (Juglans
regia) in China, with insight of Diaporthe eres complex. Mycological Progress 1–13. https://
doi.org/10.1007/s11557-018-1395-4
Gao YH, Liu F, Cai L (2016) Unravelling Diaporthe species associated with Camellia. Systemat-
ics and Biodiversity 14: 102–117. https://doi.org/10.1080/14772000.2015.1101027
Gao YH, Liu F, Duan W, Crous PW, Cai L (2017) Diaporthe is paraphyletic. IMA Fungus 8:
153–187. https://doi.org/10.5598/imafungus.2017.08.01.11
Garcia-Reyne A, López-Medrano F, Morales JM, Esteban CG, Martín I, Eraña I, Meije Y,
Lalueza A, AlastrueyIzquierdo A, RodríguezTudela JL, Aguado JM (2011) Cutaneous
infection by Phomopsis longicolla in a renal transplant recipient from Guinea: rst report of
human infection by this fungus. Transplant Infectious Disease 13: 204–207. https://doi.
org/10.1111/j.1399-3062.2010.00570.x
Gomes RR, Glienke C, Videira SIR, Lombard L, Groenewald JZ, Crous PW (2013) Diaporthe:
a genus of endophytic, saprobic and plant pathogenic fungi. Persoonia 31: 1. https://doi.
org/10.3767/003158513X666844
Guarnaccia V, Crous PW (2017) Emerging citrus diseases in Europe caused by species of Dia-
porthe. IMA Fungus 8: 317–334. https://doi.org/10.5598/imafungus.2017.08.02.07
Guarnaccia V, Crous PW (2018) Species of Diaporthe on Camellia and Citrus in the Azores
Islands. Phytopathologia Mediterranea 57(2).
Guarnaccia V, Groenewald JZ, Woodhall J, Armengol J, Cinelli T, Eichmeier A, Ezra D, Fon-
taine F, Gramaje D, Gutierrez-Aguirregabiria A, Kaliterna J, Kiss L, Larignon P, Luque J,
Mugnai L, Naor V, Raposo R, Sándor E, Váczy KZ, Crous PW (2018) Diaporthe diversity
and pathogenicity revealed from a broad survey of grapevine diseases in Europe. Persoonia
40: 135–153. https://doi.org/10.3767/persoonia.2018.40.06
Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New algo-
rithms and methods to estimate maximum-likelihood phylogenies: assessing the perfor-
mance of PhyML 3.0. Systematic Biology 59: 307–321. https://doi.org/10.1093/sysbio/
syq010
Hall T (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis pro-
gram for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.
Hu DM, Cai L, Hyde KD (2012) ree new ascomycetes from freshwater in China. Mycologia
104: 1478–1489. https://doi.org/10.3852/11-430
Huang F, Hou X, Dewdney MM, Fu Y, Chen G, Hyde KD, Li HY (2013) Diaporthe species
occurring on Citrus in China. Fungal Diversity 61: 237–250. https://doi.org/10.1007/
s13225-013-0245-6
Huang F, Udayanga D, Wang X, Hou X, Mei X, Fu Y, Hyde KD, Li HY (2015) Endophytic
Diaporthe associated with Citrus: A phylogenetic reassessment with seven new species from
China. Fungal Biology 119: 331–347. https://doi.org/10.1016/j.funbio.2015.02.006
Hyde KD, Dong Y, Phookamsak R, Jeewon R, Bhat DJ, Jones EBG, Liu NG, Abeywickrama
PD, Mapook A, Wei DP, Perera RH, Manawasinghe IS, Pem D, Bundhun D, Karunarath-
na A, Ekanayaka AH, Bao DF, Li JF, Samarakoon MC, Chaiwan N, Lin CG, Phutthacha-

Taxonomy of Diaporthe 17
roen K, Zhang SN, Senanayake IC, Goonasekara ID, ambugala KM, Phukhamsakda C,
Tennakoon DS, Jiang HB, Yang J, Zeng M, Huanraluek N, Liu JK, Wijesinghe SN, Tian
Q, Tibpromma S, Brahmanage RS, Boonmee S, Huang SK, iyagaraja V, Lu YZ, Jaya-
wardena RS, Dong W, Yang EF, Singh SK, Singh SM, Rana S, Lad SS, Anand G, Devada-
tha B, Niranjan M, Sarma VV, Liimatainen K, Aguirre-Hudson B, Niskanen T, Overall A,
Lúcio R, Alvarenga M, Gibertoni TB, Piegler WP, Horváth E, Imre A, Alves AL, da Silva
Santos AC, Tiago PV, Bulgakov TS, Wanasinghe DN, Bahkali AH, Doilom M, Elgorban
AM, Maharachchikumbura SSN, Rajeshkumar KC, Haelewaters D, Mortimer PE, Zhao
Q, Lumyong S, Xu JC, Sheng J (2020) Fungal diversity notes 115–1276: taxonomic and
phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 100:
5–277. https://doi.org/10.1007/s13225-020-00439-5
Katoh K, Toh H (2010) Parallelization of the MAFFT multiple sequence alignment program.
Bioinformatics 26: 1899–1900. https://doi.org/10.1093/bioinformatics/btq224
Kobayashi T (1970) Taxonomic studies of Japanese Diaporthaceae with special reference to
their life-histories. Bulletin of the Government Forest Experiment Station 226: 1–242.
Long H, Zhang Q, Hao YY, Shao XQ, Wei XX, Hyde KD, Wang Y, Zhao DG (2019) Dia-
porthe species in south-western China. MycoKeys 57: 113. https://doi.org/10.3897/my-
cokeys.57.35448
Mostert L, Crous PW, Kang JC, Phillips AJ (2001) Species of Phomopsis and a Libertella sp.
occurring on grapevines with specic reference to South Africa: morphological, cultural,
molecular and pathological characterization. Mycologia 93: 146–167. https://doi.org/10.
1080/00275514.2001.12061286
Murali TS, Suryanarayanan TS, Geeta R (2006) Endophytic Phomopsis species: host range and
implications for diversity estimates. Canadian Journal of Microbiology 52: 673–680. htt-
ps://doi.org/10.1139/w06-020
Rambaut A, Drummond A (2010) FigTree v.1.3.1. Institute of Evolutionary Biology, Univer-
sity of Edinburgh, Edinburgh, UK.
Rayner RW (1970) A mycological colour chart. Commonwealth Mycological Institute,
Kew, UK.
Rehner SA, Uecker FA (1994) Nuclear ribosomal internal transcribed spacer phylogeny and
host diversity in the coelomycete Phomopsis. Canadian Journal of Botany 72: 1666–1674.
https://doi.org/10.1139/b94-204
Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed
models. Bioinformatics 19: 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
Rossman AY, Farr DF, Castlebury LA (2007) A review of the phylogeny and biology of the
Diaporthales. Mycoscience 48: 135–144. https://doi.org/10.1007/S10267-007-0347-7
Santos JM, Phillips AJL (2009) Resolving the complex of Diaporthe (Phomopsis) species occur-
ring on Foeniculum vulgare in Portugal. Fungal Diversity 34: 111–125.
Santos JM, Vrandečić K, Ćosić J, Duvnjak T, Phillips AJL (2011) Resolving the Dia-
porthe species occurring on soybean in Croatia. Persoonia 27: 9–19. https://doi.
org/10.3767/003158511X603719
Udayanga D, Castlebury LA, Rossman AY, Chukeatirote E, Hyde KD (2014b) Insights into
the genus Diaporthe: phylogenetic species delimitation in the D. eres species complex. Fun-
gal Diversity 67: 203–229. https://doi.org/10.1007/s13225-014-0297-2

Qin Yang et al. / MycoKeys 67: 1–18 (2020)
18
Udayanga D, Castlebury LA, Rossman AY, Chukeatirote E, Hyde KD (2015) e Diaporthe
sojae species complex: Phylogenetic re-assessment of pathogens associated with soybean,
cucurbits and other eld crops. Fungal Biology 119: 383–407. https://doi.org/10.1016/j.
funbio.2014.10.009
Udayanga D, Castlebury LA, Rossman AY, Hyde KD (2014a) Species limits in Diaporthe: mo-
lecular re-assessment of D. citri, D. cytosporella, D. foeniculina and D. rudis. Persoonia 32:
83–101. https://doi.org/10.3767/003158514X679984
Udayanga D, Liu X, McKenzie EH, Chukeatirote E, Bahkali AH, Hyde KD (2011) e genus
Phomopsis: biology, applications, species concepts and names of common phytopathogens.
Fungal Diversity 50: 189–225. https://doi.org/10.1007/s13225-011-0126-9
Wijayawardene NN, Hyde KD, Rajeshkumar KC, Hawksworth DL, Madrid H, Kirk PM,
Braun U, Singh RV, Crous PW, Kukwa M, Lűcking R, Kurtzman CP, Yurkov A, Haelewa-
ters D, Aptroot A, Lumbsch HT, Timdal E, Ertz D, Etayo J, Phillips AJL, Groenewald JZ,
Papizadeh M, Selbmann L, Dayarathne MC, Weerakoon G, Jones EBG, Suetrong S, Tian
Q, Castanéda-Ruiz RF, Bahkali AH, Pang KL, Tanaka K, Dai DQ, Sakayaroj J, Hujslová
M, Lombard L, Shenoy BD, Suija A, Maharachchikumbura SSN, ambugala KM, Wa-
nasinghe DN, Sharma BO, Gaikwad S, Pandit G, Zucconi L, Onofri S, Egidi E, Raja HA,
Kodsueb R, Cáceres MES, Pérez-Ortega S, Fiuza PO, Monteiro JS, Vasilyeva LN, Shivas
RG, Prieto M, Wedin M, Olariaga I, Lateef AA, Agrawal Y, Fazeli SAS, Amoozegar MA,
Zhao GZ, Pfliegler WP, Sharma G, Oset M, Abdel MA, Takamatsu S, Bensch K, Silva
NI, De Kesel A, Karunarathna A, Boonmee S, Pfister DH, Lu YZ, Luo ZL, Boonyuen
N, Daranagama DA, Senanayake IC, Jayasiri SC, Samarakoon MC, Zeng XY, Doilom
M, Quijada L, Rampadarath S, Heredia G, Dissanayake AJ, Jayawardana RS, Perera PH,
Tang LZ, Phukhamsakda C, Hernández-Restrepo M, Ma XY, Tibpromma S, Gusmao LFP,
Weerahewa D, Karunarathna SC (2017) Notes for genera: Ascomycota. Fungal Diversity
86: 1–594. https://doi.org/10.1007/s13225-017-0386-0
Yang Q, Fan XL, Du Z, Tian CM (2017a) Diaporthe species occurring on Senna bicapsularis in
southern China, with descriptions of two new species. Phytotaxa 302: 145–155. https://
doi.org/10.11646/phytotaxa.302.2.4
Yang Q, Fan XL, Du Z, Tian CM (2017b) Diaporthe juglandicola sp. nov. (Diaporthales, As-
comycetes), evidenced by morphological characters and phylogenetic analysis. Mycosphere
8: 817–826. https://doi.org/10.5943/mycosphere/8/5/3
Yang Q, Fan XL, Guarnaccia V, Tian CM (2018) High diversity of Diaporthe species associated
with dieback diseases in China, with twelve new species described. MycoKeys 39: 97–149.
https://doi.org/10.3897/mycokeys.39.26914
Zhang CX, Cao ZM (2007) Primary analysis of macrofungi ora of Huoditang Mts. Journal of
Yunnan Agricultural University 22: 345–348.