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ARTICLE 3 15

/ . ) r d e g g o ) o h ng d te, i F y y an y you get f orthe oy a l sease p i d on t . 2014 . 2016). uce. l l

avoca d d Dia e t iaporthales ia scomycota emp t a axonom a v Key words: A D t phylogen h e et a ase 2015, Huan ilit . b l oeniculin '

f ens. a s ac ant pro e g l t branch cankers h f D. e y p o

8(1): 153–187 (2017) 153–187 8(1): h , ) · t t Ménard i ( D. rudis c approac aport i ces . Furthermore, species y ) Di !*#= as orthe Annesi the above conditions can be waived i l. ( f h p

ong w l y yp l http://www.indexfungorum.org . 2016 ( , cultivated worldwide in tropical introduced into new areas as reported  ) , which is a common endoph Pustulom ) MA MA al y ) a I , in Ital syn. Phomopsis ) e n t ( ogens a legalcode. Any o a / e h ca 3.0 2016 / orth ri (

p e . ions, is threatened b l ; (Gomes et a observed on carrots in France, resultin ) g m y . theicola owever, a po owever, a by-nc-nd iaporthe /

P h et a D hiodia atent pat p l ularl n. sea y . g licenses r are commonl / s e X+z ( astanea sativ

Pe (Guarnaccia C ( ( ss ytes or 2

rili h 1, e i ' ' iaporth . 2015). Because of this unique ecology and potential t l op s Mazzantia, O D d

. + y vocado as been employed for species delimitation in the genus ave been re nd subtropical re n nd rot in avocados in Chile, which was imported as been shown to cause stem and shoot cankers on sweet t a ccurately identify species of Dia ' f more than 1000 names hown to act as opportunistic plant patho hestnut urveillance, control, and trade. ole as plant pathogens, it is of paramount importance to creativecommons.org oeniculina n recent years, n seed production losses since 2007 ., indicating divergent morphological evolution. ., indicating divergent morphological evolution. // fruit from California (USA). Some endophytes have been h h i A a D of e e s f h c e r a s o + I t . la : s {'|

. t . l l d e , and Lei Ca ound at http: / f et us as ant 4,5 l s enera, namel icola et a et a t 2011, nth g i eliac

a . v g l Gomes li mptoms 2017; Published: 1 June 2017 , a e y y eo t ollowing conditions the causal V H f '“V\<V , e . n . S species are ) distinct a D 2011 y . D. an g l !*# y orthe this work, which can be mportant p f a i is the cause of p i t an icall orthe scabra h al , Van Rensburg e , Van y e crops, trees, an g 2012 p t . d Dia Pedro W. Crou Pedro W. e ª+ , iaporthe # ic D Dia n Uda 2001 (

et al vate . i l et t l l ome y % ;‚*!!!‡$]A D. vitimegaspora et a nown as

S Gomes V , k h d . ) n ng cu Grasso itis vinifera =*$$ ( %/‚X+zƒ a

di V a y ( u l % , Weijun Dua , Weijun !*! . 2011 into many smaller genera to achieve monophyly is still premature, and further collections and into many smaller genera to achieve monophyly orthe !*# * . n are p 1 . 2015 l al nc al l i sis. Several morpholo e a t t y h t e Dia tes or saprobes e in Ital e and necrosis caused b y ) et a N s parap g o Phomopsis 1 i O osts, g . Diaporthe and Stenocarpella, are embedded within aport h , O e Di z Dio s: !*# ( *, Fang Liu CTI k h / . enetic anal 2

: Submitted: 25 March 2017; Accepted: 22 Ma Submitted: 25 March 2017; ant

l l orthe neoviticol f l revious studies have shown that our understanding of species diversity within shown that our understanding of species diversity revious studies have g Ž U a 1, a i p P lo c t r D y e e ve wor : ) worldwide, and has reduced production by up to i ens, endoph Dia rom the copyright holder. Nothing in this license impairs or restricts the author’s Nothing in moral rights. rom the copyright holder. O f s or wilts on different plant hosts, several of which are plant hosts, several or wilts on different ree to share - to copy, distribute and transmit the work, under the ree to share - to copy, g . !*# ph f ME 8 · N vat mm ) y ) i U l 1 uu aport co er L ents of ' ‚H;%<\"# "‡/\+ d n- . 2011). For example, . 2011). omes pecies o owever, splitting owever, niversity of Pretoria, Pretoria 0002, South Africa 0002, South niversity of Pretoria, Pretoria O 011 l.!!` l !! Œ' g hylogenetic datasets need to be obtained to address this situation. hylogenetic datasets need to be obtained to o 2017 International Mycological Associatio 2017 International Mycological ƒ Platanus acerifolia X@]]]/'%]Hz;< State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, P.R. China P.R. Academy of Sciences, Beijing 100101, Chinese Institute of Microbiology, of Mycology, State Key Laboratory \% +z<°[‚#* !*]A/ J patho %' on many p deca et a a economically important (Mostert ‡!†U@% \ ornamentals 2 ann a G H p Article info: U a Attribution You are You INTR Yahui Gao Yahui © S * 5 1 2 # 4 Di pathogens of grapevines Abstract: TEF Phaeocytostroma <‡$‚X+zƒ; are introduced for lineages represented by multiple strains and distinct on these results, eight new species names % For any reuse or distribution, you must make clear to others the license terms o No N permission of umbel brownin been reported causing cankers and dieback on London plane was one of the most prevalent fungi isolated from grapevine perennial cankers in declining vines. ( doi:10.5598/imafungus.2017.08.01.11 V GaoG ettl al .

(Udayanga et al. 2011, Gomes et al !*# MATERIAL AND METHODS <V LE *' TEF1 !' TUB Ž# Isolates C (HIS), and calmodulin (CAL) genes are the most commonly Strains were isolated from leaves of both symptomatic and

RTI Diaporthe spp. healthy plant tissues from Yunnan, Zhejiang, and Jiangxi

A (Dissanayake et al. 2015, Udayanga et al. 2015, Huang et Provinces in China. A few other strains were obtained via the al. 2015, Santos et al. 2017). Furthermore, molecular marker +U'UE<°; aids are being used to rapidly identify Diaporthe species were isolated from imported plants from other countries. which tend to be morphologically conserved (Udayanga et Single spore isolations were conducted from diseased al !* et al. !*# “ et al !*‡ leaves with visible fungal sporulation following the protocol et al. 2015, Huang et al.!* Ž% of Zhang et al.!*#Œ boundaries remains a major challenge in Diaporthe (Huang leaf tissues was conducted following the protocol of Gao et et al. 2015), which may be a consequence of limited sampling al. (2014). Fungal endophytes were isolated according to X+z the method described by Liu et al !* Diaporthe (Liu et al. 2016). It has therefore been proposed that new species in the genus should be introduced with caution, based on cultural characteristics on PDA, spore morphology, and that multiple strains from different origins should be <V ƒ ‚' were deposited in the Mycological Herbarium, Microbiology %“et al. 2016). Institute, Chinese Academy of Sciences, Beijing, China Diaporthe with other genera in Ž@zVE'%/ Diaporthaceae Diaporthaceae General Microbiological Culture Collection Center (CGMCC). was established by Wehmeyer (1926) to accommodate Diaporthe, Mazzantia, Melanconis, and some other genera, Morphological analysis mainly based on morphological characters such as the position, Cultures were incubated on PDA at 25 °C under ambient structure, and arrangement of ascomata, stroma, and spore daylight and growth rates were measured daily for 7 d. shapes. Castlebury et al. (2002) reported that Diaporthaceae ]+z comprised Diaporthe and Mazzantia “V\ X+z (pine needle agar; Smith et al *$$` ' sequence data, removing other genera to different families %#!** /* in . Additional genera subsequently placed in incubated at a room temperature of ca. 25 oC (Su et al. 2012). the Diaporthaceae include Leucodiaporthe (Vasilyeva et al. Cultures were examined periodically for the development of 2007), Stenocarpella (Crous et al. 2006), Phaeocytostroma conidiomata and perithecia. Conidia were taken from pycnidia (Lamprecht et al. 2011), Ophiodiaporthe (Fu et al.!*# Œ Œ Pustulomyces (Dai et al. 2014). All the above genera were microscopic structures were observed and noted using a light represented by a few species or are monotypic. Although they +U"! appeared to be morphologically divergent from Diaporthe, their X

154 I MA FUNGUS Diaporthe is paraphyletic

Table 1. Sources of isolates and GenBank accession numbers used in the phylogenetic analyses of Diaporthaceae. ARTICL Species names* Culture collection Isolation sources Country GenBank Accession Numbers References no. ITS LSU TEF1 D. acaciigena /;V*$ *E' Acacia retinodes Australia ‰/#‡#!! ' ‰/#‡#_#* Gomes et al.

type) !*# E D. ampelina FAU 586 Vitiss sp. \Vz|+ ' zH‡#$`# ' ' York D. angelicae CBS 111592 Heracleum sphondylium Austria ‰/#‡#!_ ' ‰/#‡#_ # Gomes et al. !*# zA#_‡ Heracleum sphondylium Austria ‰/#‡#!` ' ‰/#‡#_ Gomes et al. !*# D. apiculata “/#‡*"E' Camellia sinensis China KP267896 KY011852 KP267970 “/#‡ Camellia sinensis China KP267901 ‰!**" # KP267975 D. arecae complex LC 4155 Rhododendron sp. China KY011895 KY011879 KY011906 LC 4159 Rhododendron sp. China KY011896 KY011880 KY011907 LC 4164 Unknown host China KY011897KY011881 KY011908 D. biguttusis “/**!`E' Lithocarpus glaber China KF576282 KY011878 KF576257 D. compacta “/#!_" Camellia sinensis China KP267850 ‰!**"#$ KP267924 “/#!"#E' Camellia sinensis China KP267854 KY011840 KP267928 “/#!"‡ Camellia sinensis China KP267855 KY011841 KP267929 D. decedens CBS 109772 Corylus avellana Austria ‰/#‡#! $ ' ‰/#‡#_" Gomes et al. !*# D. detrusa CBS 109770 Berberis vulgaris Austria ‰/#‡#!`* ' ‰/#‡#_"_ Gomes et al. !*# D. discoidispora “/# !# Camellia sinensis China KY011887KY011854 KY011898 D. elaeagni-glabrae “/‡"!E' Elaeagnus glabra China KX986779 KY011885 KX999171 LC 4806 Elaeagnus glabra China KX986780 KY011886 KX999172 D. ellipicola “/!"*!E' Lithocarpus glaber China KF576270 ‰!**"_# KF576245 D. eres “/#*$" Camellia sinensis China ‰]`_"_# KY011845 KP267947 “/#! Camellia sinensis China KP714499 KY011846 KP714511 “/#!` Camellia sinensis China KP714500 KY011847 KP714512 CBS 109767 Acer campestre Austria ‰/#‡#!_ ' ‰/#‡#"!* Gomes et al. !*# D. fusicola LC 1126 Lithocarpus glaber China KF576281 ‰!**"#` KF576256 “/!__"E' Lithocarpus glaber China ‰H _``# KY011877 ‰H _`#" D. hongkongensis LC 0784 Lithocarpus glaber China ‰/* #*!‡ KY011876 ‰/* #!$ LC 0812 Smilax china China ‰/* #*!# KY011875 ‰/* #!$‡ D. incompleta LC 6706 Camellia sinensis China ‰^$"`_$# KY011859 KX999185 “/**_E' Lithocarpus glaber China KF576267‰!**"#_ KF576242 D. mahothocarpi “/!_# Mahonia bealei China ‰/* #!$_ KY011872 ‰/* #!"" “/!_`#E' Lithocarpus glaber China ‰/* #!$` KY011871 ‰/* #!"_ D. masirevicii Diaporthe sp. Camellia sinensis China KY011888 KY011861 KY011899 D. neoarctii CBS 109490 /! \Vz|+ ‰/#‡#*‡ ' ‰/#‡#"_* Gomes et al. Jersey !*# D. oncostoma CBS 109741 Robinia pseudoacacia Russia ‰/#‡#*`* ' ‰/#‡#""_ Gomes et al. !*# D. oraccinii “/#*``E' Camellia sinensis China ‰]`_"`# ‰!**"‡# ‰]`_$#_ “/#*_ Camellia sinensis China KP267864 KY011844 ‰]`_$#" “/#$` Camellia sinensis China KP267884 KY011849 KP267958 D. ovoicicola “/**"E' Lithocarpus glaber China KF576264 ‰!**"#" ‰H _`#$ D. penetriteum “/#* Camellia sinensis China KP267879 KY011848 ‰]`_$ # “/## #E' Camellia sinensis China KP714505 KY011850 KP714517 “/##$‡ Camellia sinensis China ‰]`_"$# KY011851 KP267967 D. perjuncta CBS 109745 Ulmus glabra Austria ‰/#‡#*_ ' ‰/#‡#"$" Gomes et al. !*#

V O LUME 8 · NO . 1 155 GaoG ettl al .

Table 1. (Continued). Species names* Culture collection Isolation sources Country GenBank Accession Numbers References LE no. C ITS LSU TEF1 D. pseudophoenicicola LC 6150 Phoenix canariensis China KY011891KY011865 KY011902 RTI LC 6151 Phoenix canariensis China KY011892KY011866 ‰!**$!# A D. pustulata CBS 109742 Acer pseudoplatanus Austria ‰/#‡#*" ' ‰/#‡#$** Gomes et al. !*# CBS 109760 Acer pseudoplatanus Austria ‰/#‡#*"` ' ‰/#‡#$* Gomes et al. !*# CBS 109784 Prunus padus Austria ‰/#‡#*"_ ' ‰/#‡#$*# Gomes et al. !*# D. rudis LC 6147 Dendrobenthamia USA KY011890 KY011864KY011901 japonica LC 6145 Ilex aquifolium China KY011889 ‰!**"`# KY011900 D. saccarata /;V**`#** Protea repens, cankers South Africa ‰/#‡#*$! ' ‰/#‡#$* Gomes et al. !*# D. sclerotioides CBS 296.67 Cucumis sativus + ‰/#‡#*$# ' ‰/#‡#$*$ Gomes et al. !*# D. tectonendophytica “/``# Unknown host China KX986795 KY011857 KX999187 D. tectonigena LC 6512 Camellia sinensis China KX986782 KY011856 KX999174 D. ternstroemiae “/!___E' Ternstroemia China ‰/* #!$" KY011874 ‰/* #!"$ gymnanthera D. ueckerae “/# `‡ Camellia sinensis China KP267912 KY011855 KP267986 D. undulata LC 6624 Unknown host China KX986798 KY011858 KX999190 D. velutina LC 4414 Lithocapus sp. China KX986788 KY011882 KX999180 LC 4419 Neolitsea sp. China KX986789 ‰!**""# KX999181 “/‡‡*E' Neolitsea sp. China KX986790 KY011884 KX999182 D. xishuangbanica LC 6707 Camellia sinensis China ‰^$"`_"# KY011860 KX999175 LC 6744 Camellia sinensis China KX986784 KY011862 KX999176 D. yunnanensis LC 6168 Coffea sp. China KX986796 KY011867KX999188 Diaporthe sp. “/#* ` Camellia sinensis China KP267861 KY011842 ‰]`_$# LC 6170 Coffea sp. China ‰!**"$# KY011869 KY011904 LC 6171 Solanum melongena China KY011894 KY011870 KY011905 “/`# Theobroma cacao China KX986797 KY011868 KX999189 Mazzantia napelli zA#‡$" Aconitum vulparia Austria ' zH‡!"#`" EU222017 Castlebury et al. (2002) Ophiodiaporthe ;/A/#‡$`* Cyathea lepifera JX570889 JX570891 KC465406 Fu et al. cyatheae !*# Phaeocytostroma CPC 17071 Zea mays South Africa HA_‡"!#` ' FR748068 Lamprecht ambiguum et al. (2011) CPC 17072 Zea mays South Africa HA_‡"!#_ FR748096 FR748069 Lamprecht et al. (2011) Ph. plurivorum /;V**#"# Helianthus annuus Portugal FR748046 FR748104 FR748078 Lamprecht et al. (2011) Ph. sacchari /;V_ #‡ ' Japan FR748047 FR748105 FR748079 Lamprecht et al. (2011) Ph. megalosporum CBS 284.65 A' India FR748045 HA_‡"*!# FR748077 Lamprecht et al. (2011) Pustulomyces @H“\//**'!‡#` on dead culm of ' ‰H"!`_ # KF806755 Dai et al. bambusicola bamboo (2014) Stenocarpella CBS 117560 Rain damaged Bt maize South Africa FR748048 X°#__$#‡ ' Lamprecht macrospora !!#'!‡ et al. (2011) S. maydis CBS 117558  South Africa FR748051 X°#__$#` FR748080 Lamprecht Œ!!#!‡ et al. (2011) season Valsa ambiens CFCC 89894 Pyrus bretschneideri China KR045617 KR045699 KU710912 Fan et al. (2014) +

156 I MA FUNGUS Diaporthe is paraphyletic

(Carbone & Kohn 1999) and LR0R/LR5 primer pair (Rytas & (Jiangxi, Yunnan, Zhejiang) in the northern part of China. ARTICL Mark 1990) were used to amplify the calmodulin gene (CAL) In addition, 28 strains were isolated from the plant samples “V\ X+z % z Z U'UE < ° )“ *! & Uƒ @/2+ Bureau. {;‚ !)@+]!)@ E %! \UƒX+z The paraphyly of Diaporthe *M*!X+z]/A Phylogenetic analysis was conducted with 224 sequences parameters were as follows: 94 °C for 5 min, followed by derived from 76 ingroup taxa from Diaporthaceae with Valsa # $‡ /#! ambiens* #! /<V“V\ ` / comprised 1 817 characters including gaps (795 for LSU, 558 /z“Ž

V O LUME 8 · NO . 1 157 GaoG ettl al .

D. ellipicola LC 0810 D. eres LC 3198

LE D. biguttusis LC 1106 D. eres CBS 109767 C D. longicicola LC 1127 D. eres LC 3206 96/1 RTI D. eres LC 3205

A 99/1 D. mahothocarpus LC 0763 100/1 D. mahothocarpus LC 0732 D. penetriteum LC 3394 D. penetriteum LC 3215 D. penetriteum LC 3353

92/1 100 D. apiculata LC 3452 D. apiculata LC 3418 99/0.99 D. oraccinii LC 3172 100/1 D. oraccinii LC 3166 D. oraccinii LC 3296 100/1 D. rudis LC 6145 D. rudis LC 6147 100/1 D. arecae complex LC 4159 90/0.99 D. arecae complex LC 4155 D. pseudophoenicicola LC 6150 99/0.99 74 D. pseudophoenicicola LC 6151 71 D. arecae complex LC 4164 D. hongkongensis LC 0812 86/0.99 D. hongkongensis LC 0784 96/0.99 D. xishuangbanica LC 6744 D. xishuangbanica LC 6707 81/0.99 D. tectonigena LC 6512 D. undulata LC 6624 97/0.99 D. velutina LC 4419 100/1 D. velutina LC 4421 95/0.99 D. velutina LC 4414 100/1D. elaeagnicola LC 4802 99/1 D. elaeagnicola LC 4806 100 D. saccarata CBS 116311 D. oncostoma CBS 109741 D. decedens CBS 109772 D. ampelina FAU 586

100/1 D. angelicae CBS 111592 98/1 D. angelicae AR 3724 D. neoarctii CBS 109490 86/0.99 D. compacta LC 3078 100/1D. compacta LC 3083 72/0.99 D. compacta LC 3083 100/1 D. ueckerae LC 3564 99/1 D. tectonendophytica LC 6623 98/1 Diaporthe sp. LC 6740 Diaporthe sp. LC 6171 90/0.99 D. sclerotioides CBS 296.67 0.95 OphiOphiodiaportheiodiaporthe cyatheae BBCRCCRC 3496349611 83/0.99 0.94 Diaporthe sp. LC 6232 D. yunnanensis LC 6168 75/0.99 0.96 D. discoidispora LC 3503 Diaporthe sp. LC 6170 D. incompletap LC 6706 100/1/1PhaeocytosPPhaeocytostromastroma ambiguum** CPC 1707170711 0.97 93/1 PhaeocytosPhaeocytostromastroma ambiguum CCPCPC 1717072072 0.98 PhaeocytostromaPhaeocyto plurivorum CBS 113831138355 96/1 StenocarpellaStenoca maydis* CBS 117551175588 70 StenocStenocarpellacarpella macrospora* CCBSBS 117117560560 PhaePhaeocytostromaeocytostroma megalosporum CBS 284.65 89/1 PustulomycPustulomyces_bambusicolaces_bambusicola MFLUCC 1111_0436_0436 PhaeocytostromaPhaeocyt sacchari CCBSBS 27275.345.34 D. detrusa CBS 109770 MMazzantiaazzantia nanapellipelli AAR3498R3498 D. perjuncta CBS 109745 77 D. ovoicicola LC 1128 D. ternstroemiae LC 0777 D. fusicola LC 1126 98/0.99 Diaporthe sp. LC 3156 89 D. fusicola LC 0778 D. acaciigena CBS 129521 D. pustulata CBS 109760 87/1 D. pustulata CBS 109784 D. pustulata CBS 109742 Valsa ambiens CFCC 89894

0.07 Fig. 1. Phylogenetic tree of the family DiaporthaceaeE'<V LSU, TEF1@“%1_!†;]]1!$!% Valsa ambiens.

158 I MA FUNGUS Diaporthe is paraphyletic ARTICL E

Fig. 2. Phylogenetic tree of the genus DiaportheE'CAL, HIS, <V, TEF1, TUB@“%1_!†;]]1!$!% Diaporthella corylina.%

V O LUME 8 · NO . 1 159 GaoG ettl al . LE C RTI A

Fig. 2. (Continued).

160 I MA FUNGUS Diaporthe is paraphyletic ARTICL E

Fig. 2. (Continued).

V O LUME 8 · NO . 1 161 GaoG ettl al . LE C RTI A

Fig. 2. (Continued).

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inthif erae . u k ens ica d eb lli x r o e . vawdrey . vaccini . unshiuensis . un . uec . tu . t . t pecies names D. virgilia D. vexans D. velutina D D D D D D D D. thunbergi D D. tectonigen D. tectonend phytica D. tectona D. tecomae D. subordinaria S Table 2 Table

170 I MA FUNGUS Diaporthe is paraphyletic ) ) ARTICL 2012 ( !*# !*# !*# !*# t al. (2012 t al. . . !*# l. !*# l. !*# e e a a al al y y y y y t t t t d d d d dy dy dy ces e e e e n e r E e s stu s stu s stu s stu s stu s stu s stu f omes omes omes omes et al. omes et al. !*# omes et al. omes e his study his study his study his stud his study his study his study his study Gomes et al. Udayanga Udayanga G G Thi T T Thi T T T Thi Thi Thi Thi Thi G G T T T This study This study G G G R $ ` ‡ ` 5 6 # 1 # 4 1 2 0 9 "_ 84 7 ‡ ‡‡ ‡‡ ‡ 57 2 _ 57 5 57 ##$ # # # # 1 744 ‡ ‡ ‡ ‡ ‡ 91 $* 91 9 91 #‡#‡" 999292 999292 999 999290 19 /# /# /# /# /# / X ‰/#‡##" JX19745 J ‰ ‰ KX KY49157 KY4 KX - - - ‰^$$$" KX KX99929 ‰‡ KY4 ‰ ‰ KX KY4 KY4 - - ‰ ‰ ' CAL * $ " 0 0 # 8 "" 7 7 54 ‡ _ 2 #`#_ # #`"_ #`"_ # ‡ ‡ ‡ ‡ ‡ s #‡#_ 9992 9992 999 999266 999267 999255 r S /# /# /# /# /# / be ' ' ‰ ‰ ' ' KX KX ' ' ' ' ‰^$$$ KX KX99926 KX ‰ ‰ - - KX - KX ‰ ‰ ' HI m u * * ! # 7 4 5 9 9 0 7 5 9 8 _ 68 `# 64 62 7 5 21 5 561 5 5 n N #$ 1 o i ‡‡* ‡ ‡‡*_ ‡‡*_ ‡‡* 91 $* 91 9 91 47 7545 #‡‡* $$$#* $$$#* 999 999227 999228 999216 2 2 ^ Y49156 Y4 ^ /# /# /# / ‰/#‡‡** ‰/#‡#`# JX27545 JX ‰/# ‰/# ‰ K K ‰ KX99921 KX KX99922 ‰‡ KY4 KX KY49156 KY49157 ‰ ‰ KX KY4 KY4 KX KX99921 ‰ ‰ TUB KP ccess # $ $ " * ! 7 8 # 4 9 # 4 9 0 2 5 6 5 _ #! _ 7 14 nk A _ 55 17 55 551 55 a #" # #$ #$ #$ 1 ‡ ‡ ‡ ‡ ‡ 91 $* 91 9 91 1 754 #‡#$_ 999191 999191 999 999187 999188 9991 nB 2 /# /# /# / ‰/#‡#"_ ‰/# ‰/# KX KY49155 KY4 KX ‰^$$$*_ KX KX99918 ‰‡ KY4 KX KY49155 KY49156 ‰ ‰ KX KY4 KY4 KX KX99917 ‰ ‰ TEF Ge ' _

‡ 7 8 1 7 # 4 9 0 1 2 4 4JX # !# ! 99 99 82 95 96 4 "# 8 * ‡ 7 54 7 7 ‡ 54 7 5 7 54 _ 566 # #!!‡ # # # 1 ‡ ‡ ‡ ‡ ‡ 91 $* 91 9 91 47 577 #‡#‡ 986 986 986 986 986 $"` 2 9 X98678 X X98679 ‡ Y4 X Y49154 Y49155 X Y4 Y4 ^ X98678 F ‰/#‡#*‡ JQ619898 JX27541 J ‰/# ‰/# KX KY49154 KY4 KX K K K ‰ K K K K ‰/# ‰/# K K K ‰ K ‰/# ‰/ ITS e e y e e e aceae l aceae i nown KP aceae k eaceae ceracea ceracea ceracea etu ubiaceae ubiaceae lmaceae ost famil n olygonacea olygonacea in terculiaceae terculiaceae Roseceae P P P Corylaceae A A A B Theaceae S S - - - Fabaceae - Sterculiaceae Th R Rub R Theaceae Theaceae U Fabaceae H U e e e te yt yt y h h n n n op op e e e s s s d d g g g o o o h h h , endoph s a d i ogen s s e h ces n r e id see , ost, pat ost, pat ost, pat o in h h h ., dying stems b s sou p

rifolia sp. p., pat a n y sp., endophyte s roma cacao , en roma cacao , en lli a oro b b sp., endophyte sp., endophyte sp., endophyte abscess e ea , nown nown nown r r r cine max k k k ff m eo eo gili rus p nus y r an l amellia sinensis , endophytes n n n offea sp., endophyte offea sp., endophyte offea amellia sinensis , pathogen amellia sinensis , pathogen lmus americana y heobroma cacao , endophyt Emex australis P Emex australis Penus pentaphyll Corylus s Acer Acer Acer Al C Ca T Th Th U U U G M C Co C C C U Lupinus Isolatio Vi . o n o i E' ect # 8 ll 9 7 # .2 74 co 47

7.7 8 $`#$ 0 9 e 12 06 r 58 1

121124 ** 14 "$ 65 8 tu S S V S V MF 8108 8109 l ;V**`$ B B ; B ; MW 4 C C G / BRIP 45089b BRIP ;A<]‡ !"$ C C CGMCC 3.18291 = LC 6140 LC8113 LC8112 CGMCC 3.18292 = LC 0771 / LC 6232 LC 8114 LC 8115 L LC L L LC 6623 CGMCC 3.18289 = LC6168 LC LC 8107 CGMCC 3.18282= LC 6707 LC 6744 C / Cu C . ) * 2 1 i p. s i Continued m e sp. e ( i

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ort n e l p co aport a iaporthella corylina pecies names ab '|% P. fukushi P. P. emicis P. P. D Di Di Diaporthe sp. LC 6496 D. yunnanensis D. xishuangbanica D. woolworthi D. woodi S T

V O LUME 8 · NO . 1 171 GaoG ettl al . LE C RTI A

Fig. 3. Diaporthe acutispora /=@//#*"" A–B.#!'']+z C. Conidiomata. D–E. Conidiophores. F–G. Alpha conidia. Bars: C = 100 μm; D–G = 10 μm.

TAXONOMY Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 7.5 mm diam/d. Colony entirely white Diaporthe acutispora Y.H. Gao & L. Cai, sp. nov. %\ MycoBank MB820679 aerial mycelium. H# Additional material examined: China: Yunnan Province: Xishuangbanna, Etymology|+. on healthy leaves of Camellia sasanqua, 20 Sep. 2014, W.J. Duan, culture LC 6142; ibid. culture LC 6160. Diagnosis: Diaporthe acutispora is phylogenetically distinct and morphologically differs from species reported from the Diaporthe elaeagni-glabrae Y.H. Gao & L. Cai, sp. nov. host genera Coffea and Camellia in the larger conidiophores MycoBank MB820680 # (Fig. 4)

Type: China: Yunnan Province: Aini Farm, on healthy leaves of Etymology: +Elaeagnus glabra. Coffea sp., 20 Sep. 2014, W.J. Duan (HMAS 247086 – holotype, {/=@//#*"" —“/`*`*ME' Diagnosis: Diaporthe elaeagni-glabrae can be distinguished from the closely related species D. elaeagni $` † <V Description| „ ]+z| Conidiomata pycnidial, globose, $# † TEF1$‡ † TUB$` † HIS$‡ †CAL) $$M‡_# and D. stictica $` †<V$ † TEF$_ †TUB$` † μm diam, often with a yellowish conidial cirrus exuding from in HIS$` † CAL) (Fig. 2). Diaporthe elaeagni-glabrae the ostioles. Conidiophores *!M#‡ & M# 3 differs from other species recorded from Elaeagnus in the hyaline, septate, branched, straight or slightly curved, # tapering towards the apex. Alpha conidia abundant in culture, _M*! &M#3 x —"‡/!_&`/! n—#! Type: China: Jiangxi Province: on diseased leaves of Elaeagnus ' Beta conidia not glabra V!*# Y.H. Gao (HMAS 247089 – holotype, dried observed. {/=@//#*""_—“/‡"!ME'

172 I MA FUNGUS Diaporthe is paraphyletic ARTICL ) ) ) ) ) ) 2014b 2014c 2014a ( ( ( ) ) )

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y y g o a omes ao a ao ao uang efe G G G Ud Chang ] Uda Diedicke Petch Uecker Uecker Udayanga G G H Uecker (1988) Dio R M *#M : working collection of Pedro Crous C **M P *‡M*` C : Working collection o LC : Working & **M ! `

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f c i an ora a , CAS, China; MFL p i a acea gb l i l lin leta ii ora teum ae gy a i r p n p i acta co cu cu s ii p gdal tin uan eae ni ni y u oldiae h l es ff s aeagn r enetr e i unnanensis l rn oe oe heae e p uecke y ecie a co China General Microbiological Culture Collection; . elaeagni-glabrae . incom . v . oracc . . . x . apiculata . com . discoidis : p . acac . t . . e D D D D D D D P P P P D. D. f D. f D. hongkongensi D. elaeagnicola P. D D D S D. am |<V@@“\VXz'zAV;%@\Vz{ ynoptic characters o s S FAU . , CGMCC nu 3 a g sea

e lit DP l , aea eo hina. offea P. ab : Australian plant pathogen culture collection, Queensland, Australia; Australian plant pathogen culture collection, Queensland, BRIP : at Institute of Microbiolo C N AR maintained at the Westerdijk Institute maintained at the Westerdijk El gener Host C T Camellia D. acutis

V O LUME 8 · NO . 1 173 GaoG ettl al . LE C RTI A

Fig. 4. Diaporthe elaeagni-glabrae/=@//#*""_A–B.*‡'']Xz{ C. Conidiomata; D–H. Conidiophores; I. Alpha conidia; J. Beta conidia. Bars: C = 100 μm; D–J = 10 μm.

Description: On PDA: Conidiomata ##!M**_! erumpent or immersed in medium, translucent conidia μm, erumpent, with slightly elongated black necks, yellowish E**!M#"!3 Conidiophores or dirty white, spiral conidial cirri extruding from ostioles. cylindrical, straight or sinuous, apical or base sometimes Conidiophores *`M" & * M 3 ** M# &*"M# 3 x —*`/#& /! \Alpha conidia`M*#& —#!Beta conidia% * M#3 x —"#/*‡&/!# n—#! E** M#&! M3 x or oval, usually biguttulate. Beta conidia_ M &*M3 —!/_ &*/!‡—!Alpha conidia not observed. x —* */# &*/! n—‡! curved, base truncate. Culture characters: Cultures on PDA at 25 °C in dark, with 12/12 h alternation between daylight and darkness pure Culture characters: Cultures incubated on PDA at 25 °C in white (surface) and pale yellow to cream (reverse). Colony darkness, growth rate 7 mm diam/d. Colony pale yellowish, pellicular, forming less pigmented sectors, with concentric greenish to brownish at the centre, reverse pale yellowish rings of gummy mycelium. Growth rate was 10.5 mm diam/d. and brownish at the centre with age. Aerial mycelium white, \ % Material examined: Ukraine: from seeds of Helianthus annuus#! at maturity. Oct. 2015, W.J. Duan “/ `*_# M Japan: Lagerstroemia indica#!„!* W.J. Duan, culture LC 6185. Additional material examined: China: Jiangxi Province: on diseased leaves of Elaeagnus glabra V!*# Y.H. Gao, culture LC Notes: Diaporthe helianthi, responsible for stem canker 4806. \ Helianthus annuus) @'/%% et al. 1981), has been listed in the Diaporthe helianthi @'/%et al., Nova Hedwigia / ƒ % 34|‡##*$"* \ ƒ (Fig. 5) \ Description: Sexual morph not produced. Conidiomata seeds from Ukraine and Lagerstroemia indica from Japan. pycnidial globose to subglobose, dark brownish to black,

174 I MA FUNGUS Diaporthe is paraphyletic ARTICL E

Fig. 5. Diaporthe helianthi (LC 6185). A–B. _'']Xz{C. Conidiomata; D–F. Conidiophores; G–H. Beta conidia. Bars: C = 100 μm; D–H = 10 μm.

Diaporthe incompleta Y.H. Gao & L. Cai, sp. nov. conidial droplets exuding from the central ostioles. MycoBank MB820681 Conidiophores"M&*M 3 (Fig. 6) unbranched, smooth, slightly curved, tapering towards apex. Alpha conidia not observed. Beta conidia *$M‡‡ & ! M* Etymology|+. μm ( x —#! /"_&**/!‡ n—#! % Diagnosis: Diaporthe incompleta is phylogenetically distinct and differs morphologically from other species recorded from Culture characters: Cultures incubated on PDA at 25 °C in Elaeagnus and Camellia in the much longer beta conidia darkness, growth rate 16.5 mm diam/d. Colony entirely white, # \ % Œ the centre with age. Aerial mycelium white, cottony, margin Type: China: Yunnan Province: Xishuangbanna, on diseased of lobate, conidiomata visible at maturity. Elaeagnus glabra, 19 Apr. 2015, F. Liu (HMAS 247088 – holotype, {/=@//#*"""—“/`_ ‡ME' Additional material examined: China: Yunnan Province: Xishuangbanna, on diseased leaves of Camellia sinensis, 19 Apr. Description: Conidiomata pycnidial, subglobose to globose, 2015, F. Liu, culture LC 6706. brownish to black, 207–650 μm diam, cream to pale luteous

V O LUME 8 · NO . 1 175 GaoG ettl al . LE C RTI A

Fig. 6. Diaporthe incompleta /=@//#*"""A. Leaves of host plant; B–C._''{D. Conidiomata; E–F. Conidiophores; G. Beta conidia. Bars: D = 100 μm; E–G = 10 μm.

Diaporthe podocarpi-macrophylli Y.H. Gao & L. Cai, Diagnosis: Diaporthe podocarpi-macrophylli can be distin' sp. nov. guished from the phylogenetically closely related species D. MycoBank MB820682 pseudophoenicicola$_†<V$!†TEF1, 98 (Fig. 7) † TUB$_† HIS$_† CAL). Morphologically, D. podocarpi-macrophyllii differs from other species occurring Etymology: + Podocarpus on the host genera Podocarpuss and Olea, i.e. D. cinerascens macrophyllus. and Phomopsis podocarpii in its wider and shorter alpha co'

176 I MA FUNGUS Diaporthe is paraphyletic ARTICL E

Fig. 7. Diaporthe podocarpi-macrophylli /=@//#*""* A–B.#!'']Xz{C. Conidiomata; D–F. Conidiophores; G–I. Alpha and beta conidia. Bars: C = 100 μm; D–I = 10 μm.

nidia and the presence of beta conidia (Chang et al. 2005, *! M_&* M 3 x —* #/‡#&*/!# n—#! Gomes et al.!*#{|'% cylindrical to clavate, hyaline, septate, branched, smooth, straight. Alpha conidia# M" &*M#3 x —`#/*_&* Type: Japan: on healthy leaves of Podocarpus macrophyllus, ± 0.7, n = 50), unicellular, aseptate, fusiform, hyaline, usually 20 Sep. 2014, W.J. Duan (HMAS 247084 – holotype, dried biguttulate and acute at both ends. Beta conidia" M#* & {/=@//#*""*—“/`* ME' 0.5–2 μm ( x — *$ / _* & ** / !‡ n — #! % Description: Conidiomata pycnidial in culture on PDA, solitary ends, base truncate. or aggregated, deeply embedded in the PDA, erumpent, dark brown to black, 222–699 μm diam, yellowish translucent Culture characters: Cultures incubated on PDA at 25 °C in conidial drops exuding from the ostioles. Alpha conidiophores * / `M*"&* M#3 x —*#/`&*/!# n—#! \ septate, branched, cylindrical, straight to sinuous, sometimes mycelium, reverse pale brown with brownish dots with age, \ Beta conidiophores with visible solitary or aggregated conidiomata at maturity.

V O LUME 8 · NO . 1 177 GaoG ettl al . LE C RTI A

Fig. 8. Diaporthe undulata/=@//#*"$#A. Leaves of host plant; B–C.#!'']+z{D. Conidiomata; E. Conidiophores; F–G. Alpha conidia. Bars: D = 100 μm; E–G = 10 μm.

Additional material examined: Japan: on healthy leaves of Description: Conidiomata pycnidial, irregular, embedded in Podocarpus macrophyllus, 20 Sep. 2014, W.J. Duan, culture "M ‡#3 LC 6141; ibid. culture LC 6144; ibid. culture LC 6156; ibid. culture with short hyphae, one to several necks forming from a single LC 6157. – China: Zhejiang Province: on healthy leaves of P. pycnidium. Conidiophores obpyriform, hyaline, phiailidic, macrophyllus, 10 Jul. 2015, W.J. Duan, culture LC 6194; ibid. M*_ &M#3 x —$_/‡!&‡/ culture LC 6195; ibid. culture LC 6196; ibid. culture LC 6197; ibid. 0.5, n = 20). Alpha conidia ellipsoid, hyaline, biguttulate, culture LC 6198; ibid. culture LC 6199; ibid. culture LC 6200; ibid. M` &M# x — "/!‡&#/!# culture LC 6201; ibid. culture LC 6202; ibid.“/`# MItaly: n = 50). Beta conidia not observed. on healthy leaves of Olea europaea, 20 Sep. 2014, W.J. Duan, culture LC 6229. Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 10.5 mm diam/d. Colony entirely white, Diaporthe undulata Y.H. Gao & L. Cai, sp. nov. reverse pale yellowish and dark brownish at the centre with @;@;"!`"# age. Aerial mycelium white, cottony, dense, with undulate (Fig. 8) margin and visible conidiomata at maturity.

Etymology: +R Additional material examined: China-Laos border: unknown host, 19 Apr. 2014, F. Liu, culture LC 8110; ibid. culture LC 8111. Diagnosis: Diaporthe undulata differs from the most closely related species, D. biconispora,%$‡†<V Diaporthe velutina Y.H. Gao & L. Cai, sp. nov. "‡† TEF1$#† TUB), and from otherr Diaporthe MycoBank MB820684 species in the obpyriform conidiophores and shorter and (Fig. 9) # Etymology|+ felted colony. Type: China-Laos border: on diseased leaves of unknown host, 19 Apr. 2014, F. Liu (HMAS 247091 – holotype, dried Diagnosis: Diaporthe velutina is distinguished from D. {/=@//#*"$#—“/``‡ME' anacardii<VTEF1, TUBB and HISS$$†<V $ † TEF1 $$ † TUB $" † HIS), and from

178 I MA FUNGUS Diaporthe is paraphyletic ARTICL E

Fig. 9. Diaporthe velutina /=@//#*""`A. Diseased leaves; B–C. #!'']Xz{D. Conidiomata; E. Conidiophores; E. Alpha and beta conidia. Bars: D = 100 μm; E–F = 10 μm.

other Diaporthe species reported from Camellia sinensis in dense, felted, conidiomata erumpent at maturity, reverse %Œ# centre yellowish to brownish.

Type: China: Jiangxi Province: on diseased leaves of Neolitsea Additional material examined: China: Jiangxi Province: Yangling, on V!*# Y.H. Gao (HMAS 247087 – holotype, dried diseased leaves of Neolitsea V!*#Y.H. Gao, culture {/=@//#*""`—“/‡‡*ME' LC 4419; ibid.“/‡‡{=+\% #z!*# Q. Chen, culture LC 4788; Fengshan, on diseased Description: Conidiomata pycnidial, globose, black, embed' leaves of Callerya cinerea V!*#Y.H. Gao, culture LC 4641. ded in PDA, aggregated in clusters, 69–428 μm diam, cream Yunnan Province: Xishuangbanna, on diseased leaves of Camellia translucent drop of conidia exuded from the central ostioles. sinensis, 19 Apr. 2015, F. Liu, culture LC 6708; loc. cit., on healthy Conidiophores *!M#&*M 3 leaves of C. sinensis, 21 Apr. 2015, F. Liu, culture LC 6519. branched, densely aggregated, slightly tapering towards the apex, sometimes slightly curved. Alpha conidia M*! & Diaporthe xishuangbanica Y.H. Gao & L. Cai, sp. 2–2.5 μm ( x —`$/!$&/! n = 50), unicellular, nov. %' MycoBank MB820685 ' Beta conidia**M_ &! M* 3 x = 16.1 (Fig. 10) / !&!"/!‡ n — #! E rounded, curved. Etymology|+^

Culture characters: Cultures incubated on PDA at 25 °C in Diagnosis: Diaporthe xishuangbanica can be distinguished darkness, growth rate 18.75 mm diam/d. Colony entirely from the phylogenetically closely related D. tectonigena in white, surface mycelium greyish to brownish at the centre, %$"†<V$!† TEF1$`†TUB)

V O LUME 8 · NO . 1 179 GaoG ettl al . LE C RTI A

Fig. 10. Diaporthexishuangbanica/=@//#*""# A–B._'']Xz{ C–D. #!'']+z{E. Conidiomata; F–K. Conidiophores; L–N.z;|U—*!!3{HM+—*!3

(Fig. 2), and from other Diaporthe species reported from Diaporthe yunnanensis Y.H. Gao & L. Cai, sp. nov. Camellia in the longer conidiophores and alpha conidia MycoBank MB820686 # (Fig. 11)

Type: China: Yunnan Province: Xishuangbanna, on dis' Etymology|+ eased leaves of Camellia sinensis, 19 Apr. 2015, F. Liu collected, Yunnan Province. Ž@zV‡_!"#M{/=@//#*""#— “/`_‡‡ME' Diagnosis: Diaporthe yunnanensiss can be distinguished from the phylogenetically closely related D. siamensis $` † Description: Conidiomata *"!M#*! 3 <V$*†TEF1$‡† TUB) (Fig. 2), and from other diam, scattered on the pine needle. Conidiophores cylindrical, Diaporthe species reported on the genus Camellia in the *#M#‡ &* M#3 x —!$/ &*/!# n = 40), # branched, septate, straight, sometimes sinuous or lateral. Alpha conidia_M$ & M# 3 x —"#/!_&"/!# Type: China: Yunnan Province: Xishuangbanna, on healthy n — #! ' Beta conidia not leaves of Coffea sp., 20 Sep. 2014, W.J. Duan (HMAS observed. ‡_!$` M { /=@// #*""$ — “/ `*`"ME' Culture characters: Cultures incubated on PDA at 25 °C in darkness, growth rate 17.5 mm diam/d. Colony entirely Description: Conidiomata pycnidial, 195–880 μm diam, white, reverse pale yellowish to greenish. Aerial mycelium globose or irregular, erumpent, solitary or aggregated %%% together, dark brown to black. Conidia exuding from the at maturity. pycnidia in white to cream drops. Conidiophores cylindrical, straight or slightly curved. Alpha conidia#M` &*M 3 Additional material examined: China: Yunnan Province: Xishuangbanna, x — /*&/! n—#! on diseased leaves of Camellia sinensis, 19 Apr. 2015, F. Liu, culture LC with one end obtuse and the other acute. Beta conidia*# M `_!_/=@//#*"" ## &*M* 3 x —_ / &* /!#n—#!

180 I MA FUNGUS Diaporthe is paraphyletic ARTICL E

Fig. 11. Diaporthe yunnanensiss/=@//#*""$ A–B._'']Xz{C. Conidiomata; D. Conidiophores; E. Alpha and beta conidia; F. Beta conidia. Bars: C = 100 μm; D–F = 10 μm.

aseptate, hamate or curved, base truncate. observed, base subtruncate. Beta conidia* M‡! &*M μm ( x —`!/ "&*"/! n—#! Culture characters: / ]Xz \ curved, base subtruncate, tapering towards one apex. growth rate of 5.5 mm diam/d, with abundant dirty white and yellowish pigmented mycelium, dry, felted, extensive thin, Culture characters: Cultures incubated on PDA at 25 °C in and in reverse the centre cream, with zone rings of pale to _"#  / dark brownish pigmentation. white to dirty pink, cottony, sparse, brownish to black conidiomata erumpent at maturity, coated with white hypha, Additional material examined: China: Yunnan Province: Xishuangbanna, granular at margin, reverse pale brown, with brownish dots on healthy leaves of Coffea sp., 20 Sep. 2014, W.J. Duan, culture LC when maturity. 8106; ibid. culture LC 8107. Material examined: China: Zhejiang Province| = + Diaporthe sp. 1 A%$:!+*":*‡U % Alnus mill, Jan. 2010, (Fig. 12) Y.Y. Suu/=@//#*"$—“/!__*

Description: Conidiomata pycnidial, subglobose to globose, Notes| dark brown to black, deeply embedded in the substrate, complex, which is reported from a very wide range of host scattered on the substrate surface, embedded in PDA, plants and includes mostly opportunistic pathogens or clusters in group of 2–7 pycnidia, 268–509 μm, yellowish secondary invaders on saprobic host substrata (Udayanga drop of conidia diffusing from the central ostioles. et al. 2014a, Gao et al. 2016). Species delimitation in this Conidiophores ` M*$ & *M# 3 complex is currently unclear. Udayanga et al. (2015) accepted \ nine phylogenetic species in the D. eress complex, including tapering towards the apex. Alpha conidia_ M*# &M# D. alleghaniensis, D. alnea, D. bicincta, D. celastrina, D. μm ( x —$$/*‡&"/!‡n—#! eres, D. helicis, D. neilliae, D. pulla, and D. vaccinia. Gao fusoid to ellipsoid or clavate, two or several large guttulae et al. (2016) examined 17 isolates belonging to the D. eres

V O LUME 8 · NO . 1 181 GaoG ettl al . LE C RTI A

Fig. 12. Diaporthe*/=@//#*"$ A. Leaves of host plant; B–C. #!'']Xz{D. Conidiomata; E–F. Conidiophores; G. Beta conidia; H–I. Alpha conidia. Bars: D = 100 μm; E–I = 10 μm.

complex, and reported that many presented intermediate E morphology among “species” and the phylogenetic analyses group remains unresolved. often resulted in ambiguous clades with short branch and

182 I MA FUNGUS Diaporthe is paraphyletic

Diaporthe sp. 2 Diaporthe eucommiicola (C.Q. Chang et al.) Y.H. ARTICL Gao & L. Cai, comb. nov. Culture characters: Cultures incubated on PDA at 25 °C in MycoBank MB821441 #"#  / Basionym: Phomopsis eucommiicola C.Q. Chang et al., My- convex, entire white to yellowish, reverse brownish. Aerial cosystema 24: 147 (2005). E ' Type: China: Hunan Province: on living branches of Material examined: Japan: on leaves of Acer sp., 20 Sep. 2014, W.J. Duan, Eucommia ulmoidess and Styrax hypoglauca, L.J. Luo (SCHM /=@//#*"$*—“/`*‡!“/"**{ ibid.“/"**# #`!_M{z _"!_*<Vƒ% V/Ž@#`!_ Notes: Although three isolates clustered in a clade distinctly different from known species in the genus included, they are Diaporthe glabrae (C.Q. Chang et al.) Y.H. Gao & L. not formally described because they were sterile. Diaporthe Cai, comb. nov. sp. 2 shares a low homology to the most closely related @;@;"*‡‡# species, D. rhoina$ †<V"_†TEF1$_†TUB, Basionym: Phomopsis glabrae C.Q. Chang et al., My- $‡ † HIS $ † CAL). Five Diaporthe species cosystema 24: 8 (2005). are so far only known from the sterile state, including D. endophytica, D. inconspicua, D. infecunda, D. asheicola, and Type: China: Fujian Province: on living branches of Bougainvillea D. sterilis (Gomes et al. !*#“et al. 2014). glabra, Y.H. Chengg V/Ž@ #` M {z`!*$*" <V ƒ%V/Ž@#` Diaporthe averrhoae (C.Q. Chang et al.) Y.H. Gao & L. Cai, comb. nov. Diaporthe lagerstroemiae (C.Q. Chang et al.) Y.H. @;@;"*‡#_ Gao & L. Cai, comb. nov. Basionym: Phomopsis averrhoae C.Q. Chang et al., My- MycoBank MB821444 cosystema 24: 6 (2005). Basionym: Phomopsis lagerstroemiae C.Q. Chang et al., My- cosystema 24: 148 (2005). Type: China: Fujian Province: on living branches of Averrhoa carambola, Y.H. ChenggV/Ž@#`! M{z`*"$#! Type: China: Hunan Province: on living branches of <Vƒ%V/Ž@#`! Lagerstroemia indica, C.Q. ChanggV/Ž@#`!"M{ z`$$‡<Vƒ%V/Ž@ Diaporthe camptothecae (C.Q. Chang et al.) Y.H. #`!" Gao & L. Cai, comb. nov. @;@;"*‡#" Diaporthe liquidambaris (C.Q. Chang et al.) Y.H. Gao Basionym: Phomopsis camptothecae C.Q. Chang et al., Myy- & L. Cai, comb. nov. cosystema 24: 145 (2005). MycoBank MB821446 Basionym: Phomopsis liquidambaris C.Q. Chang et al., My- Type: China: Hunan Province: on living branches of Camp- cosystema 24: 9 (2005). totheca acuminate, L.J. Luo V/Ž@ #`** M { z`$$`<Vƒ%V/Ž@ Type: China: Fujian Province: on living branches of Liquii- #`** dambar formosana, Y.H. Chengg V/Ž@ #`* M { z`!*$*$<Vƒ%V/Ž@ Diaporthe chimonanthi (C.Q. Chang et al.) Y.H. Gao #`* & L. Cai, comb. nov. @;@;"*‡#$ Diaporthe loropetali (C.Q. Chang et al.) Y.H. Gao & L. Basionym: Phomopsis chimonanthi C.Q. Chang et al., My- Cai, comb. nov. cosystema 24: 146 (2005). MycoBank MB821448 Basionym: Phomopsis loropetalii C.Q. Chang et al., My- Type: China: Hunan Province: on living branches of Chimonanthus cosystema 24: 148 (2005). praecox, C.Q. ChanggV/Ž@#`*‡M{z`$$#<V ƒ%V/Ž@#`*‡ Type: China: Hunan Province: on living branches of Loropetalum chinense, C.Q. ChanggV/Ž@#`* M{ Diaporthe eucommiae (F.X. Cao et al.) Y.H. Gao & L. z`!*$*_<Vƒ%V/Ž@ Cai, comb. nov. #`* MycoBank MB821440 Basionym: Phomopsis eucommiae F.X. Cao et al., J. Middle- Diaporthe magnoliicola Y.H. Gao & L. Cai, nom. nov. South China Forestry Coll. 10|#‡*$$!{eucommi’. MycoBank MB821459 Replaced name: Phomopsis magnoliae M.M. Xiang et al., Type: China: Guangdong Province: from leaves of Eucommia My-cosystema 21: 501 (2002). ulmoides, F.X. CaoV/Ž@!!!M{z`!*$*<V sequence derived from the holotype SCHM 0020).

V O LUME 8 · NO . 1 183 GaoG ettl al .

Type: China: Guangdong Province: on leaves of Magnolia Fu et al.!*#X et al. 2014). For example, Ophiodiaporthe coco, Z.D. Jiang V/Ž@ #!!* M {z`$$ <V produces only one type of globose or subglobose conidia LE ƒ%V/Ž@#!!* C conidia of Diaporthe (Fu et al!*#{ Phaeocytostroma and

RTI Note| magnoliae is occupied, so Diaporthe Stenocarpella produce pigmented alpha conidia which differ

A magnoliicola is proposed as a replacement name. from the hyaline conidia of Diaporthe (Lamprecht et al. 2011); Pustulomyces produces larger, straight or sigmoid conidia Diaporthe michelina (C.Q. Chang et al.) Y.H. Gao & L. (Dai et al. 2014). Phaeocytostroma and Stenocarpella were Cai, comb. nov. originally suspected to be members of Botryosphaeriaceae MycoBank MB821460 (Botryosphaeriales) because of their pigmented alpha conidia Basionym: Phomopsis michelina C.Q. Chang et al., My- '/ et al. 2006). However, cosystema 24: 9 (2005); as ‘micheliae’. they were subsequently allocated to Diaporthales based on phylogenetic analysis (Lamprecht et al. 2011), which is Type: China: Fujian Province: on living branches of Michelia alba, Y.H. Chengg V/Ž@ #`!# M {z`!"!<V ODiaporthe” clade embedded with the hetero' ƒ%V/Ž@#`!# geneous genera Ophiodiaporthe, Pustulomyces, Phaeocyy- tostroma, and Stenocarpella is probably a typical example Diaporthe phyllanthicola (C.Q. Chang et al.) Y.H. of divergent evolution in morphological characters. Such an Gao & L. Cai, comb. nov. evolution could have been driven by host and/or environmen' MycoBank MB821461 tal adaptations. For example, the monotypic Ophiodiaporthe Basionym: Phomopsis phyllanthicola C.Q. Chang et al., My- is associated with Cyathea lepifera (a fern), while Pustulomyy- cosystema 24: 10 (2005). cess is bambusicolous (Dai et al. 2014). On the contrary, none of the previously named over 1 900 Diaporthe / Phomopsis Type: China: Fujian Province: on living branches of species was recorded from a fern or Bambusaceae (https:// Phyllanthus emblica, Y.H. Chengg V/Ž@ #`"! M { '% < z`!"*$<Vƒ%V/Ž@ to speculate that the speciation of Ophiodiaporthe and Pustu- #`"! lomyces, as well as the distinctly different morphologies from their close Diaporthe allies, are the consequences of evo' lutionary adaption to new hosts. Similarly, Phaeocytostroma DISCUSSION and Stenocarpella are mainly restricted to maize (Zea mays), causing root stalk and cob rot (Stovold et al. 1996, Lamprecht In this study, eight new species of Diaporthe are introduced, et al. 2011). having been isolated from various plant hosts collected in Splitting Diaporthe into many smaller genera would %Phomopsis species described from achieve monophyletic groupings, but would also create many China were subjected to molecular analysis, and transferred OPDiaporthe to Diaporthe to conform to the “one fungus one name” rule would have no recognisable morphological distinctions (Udayanga et al. 2011, Rossman et al!*` in either sexual or asexual morphs. In addition, splitting of the other Phomopsiss species described from Diaporthe into many smaller genera will result in numerous /'% name changes, which is certainly an unfavourable option for Diaporthe. both mycologists and plant pathologists. Previous taxonomic studies in Diaporthe (syn. Phomopsis) Diaporthe ' have been primarily based on morphology, which has been pathogens, some on economically important hosts, such as \ % Helianthus annuus \{ et al. 2011) and due to the simple and plastic morphological characters Glycine maxx (; Santos et al. 2011). However, the (Gao et al!* number of known endophytic Diaporthe species has increased of ascomycetes. For example, species referred to Phoma rapidly in recent years (Huang et al. 2015, Gao et al. 2016). have been shown to be highly polyphyletic and scattered Wang et al. !*# throughout at least six families within Pleosporales (Aveskamp ecology and biology of endophyticc Diaporthe species is just the et al. 2010, Chen et al. 2015). Although Diaporthe was OP<!*# previously thought to be monophyletic based on its typical Diaporthe endophytica, was formally named (Gomes et al. and unique Phomopsiss asexual morph and diaporthalean !*#Citruss conducted by Huang et al. (2015) sexual morph (Gomes et al. !*# recorded seven apparently undescribed endophytic Diaporthe is revealed in the present study (Fig. 1). Several genera, species. Inspection of Diaporthe species on Camellia sinensis notably Ophiodiaporthe (Fu et al. !*# Pustulomyces % (Dai et al. 2014), Phaeocytostroma, and Stenocarpella all occurring as endophytes (Gao et al. 2016). Because many (Lamprecht et al. 2011), are shown to be embedded in of these plant pathogenic Diaporthe species are commonly Diaporthe s. lat., none of which present an independent X+z lineage from Diaporthe as currently circumscribed (Fig. 1). z characteristics (Vasilyeva et al. 2007, Lamprecht et al. 2011, of quarantine and disease control (Udayanga et al. 2011).

184 I MA FUNGUS Diaporthe is paraphyletic

et al. !** lichenized, and other fungi. Plant Systematics and Evolution 216: ARTICL \ z Diaporthe helianthii which ‡#M‡$ was originally only known from Europe (former Yugoslavia), X X° J‚ ++ ; XZ / U ; and is apparently an invasive species in Australia. Diaporthe AH, et al. (2014) Pustulomyces gen. nov. accommodated in helianthii is listed in the Chinese quarantine directory, and Diaporthaceae, Diaporthales, as revealed by morphology and E has long been considered a predominant disease limiting molecular analyses. Cryptogamie, Mycologie 35|`#M_ production in Europe (Desanlis et al. !*# X et al. XX=“XA]X!*‚@| !*`\ more models, new heuristics and parallel computing. Nature from Ukraine into China. Here, we report another interception Methods 9: 772. of D. helianthi from Lagerstroemia indica imported from X @z Z+ @ U X ] !*#z Z/%% \\Phomopsis helianthi quickly serious pathogens such as Diaporthe species can epidemics as a function of cropping practices. Field Crops be distributed as endophytes or latent pathogens with global Research 149|`#M_ trade. Diogo EL, Santos JM, Phillips AJ (2010) Phylogeny, morphology and pathogenicity of Diaporthe and Phomopsiss species on almond in Portugal. Fungal Diversityy 44: 107–115. ACKNOWLEDGEMENTS Dissanayake AJ, Liu M, Zhang W, Chen Z, Udayanga D, et al. (2015) Morphological and molecular characterisation of Diaporthe J“/R species associated with grapevine trunk disease in China. + + V Fungal Biologyy 119|"#M$‡ H / +VH/ #***!*!#$!` @ X@XzZJX+;V“Z‰ V/@„V!*‡H*!*!! et al. (2017) Microfungi on Tectona grandis (teak) in northern Fungal Diversity 82: 107–182. Du Z, Fan XL, Hyde KD, Yang Q, Liang YM, et al. (2016). Phylogeny REFERENCES and morphology reveal two new species of Diaporthe from Betula spp. in China. Phytotaxa 269: 90–102. z “ “ ª V = @ ; z !* X JZ X “Z / ^H / “ !*` < Characterization and pathogenicity of Phomopsis theicola quarantine fungus Diaporthe helianthi from the corn seeds anamorph off Diaporthe foeniculina causing stem and shoot imported from Ukraine. Mycosystema 35|* !#M* *# cankers on sweet chestnut in Italy. Journal of Phytopathology H^“Ž‰X\XJ^/@!* Diaporthe 164: 412–416. rostrata, a novel ascomycete from Juglans mandshurica Aveskamp MM, de Gruyter J, Woudenberg JHC, Verkley GJM, associated with walnut dieback. Mycological Progress 14: 82. Crous PW (2010) Highlights of the Didymellaceae: a polyphasic H^“/@°“@/Z et al. (2014) Cytospora approach to characterise Phoma and related pleosporalean from Salixx in northern China. Mycotaxon 129|#!#M#* genera. Studies in Mycology 65: 1–60. H /Ž Ž Ž@ / // Ž@ et al. !*# Carbone I, Kohn LM (1999) A method for designing primer sets for Ophiodiaporthe cyatheae gen. et sp. nov., a diaporthalean Mycologia 91: pathogen causing a devastating wilt disease of Cyathea lepifera #M ` Mycologia 105: 861–872. Castlebury LA, Rossman AY, Jaklitsch WJ, Vasilyeva L (2002) A =ŽVJV/“!*‡Phomopsis preliminary overview of the Diaporthaless based on large subunit in Gutianshan nature reserve in China. Mycological Progress 13: X+zƒMycologia 94|*!*_M*!#* 111–121. //°/Ž^@@Z[X!! + Gao YH, Su YY, Sun W, Cai L (2015) Diaporthe species occurring Phomopsis on woody plants in Fujian Province. Mycosystema on Lithocarpus glabra / % 24: 6–11. species. Fungal Biology 119|$ M#!$ Chen Q, Jiang JR, Zhang GZ, Cai L, Crous PW (2015) Resolving the Gao YH, Liu F, Cai L (2016) Unravelling Diaporthe species associated Phoma enigma. Studies in Mycology 82|*#_M*_ with Camellia. Systematics and Biodiversity 14: 102–117. Chi PK, Jiang ZD, Xiang MM (2007) Flora Fungorum Sinicorum. Vol. = +“ X =/ *$$ X% #‡ Phomopsis. Beijing: Science Press. designed for use with the PCR to amplify conserved genes /]J=Z[AZ@V]Ž'Z Applied and Environmental +“ !!‡ Calonectria species and their Cylindrocladium Microbiology 61|*##M*##! anamorphs: species with sphaeropedunculate vesicles. Studies Gomes R, Glienke C, Videira S, Lombard L, Groenewald J, et al. in Mycology 50|‡* M‡#! !*# Diaporthe: a genus of endophytic, saprobic and plant / ]J V ; J @Z A Z @ JH„ pathogenic fungi. Persoonia 31: 1–41. et al. (2006) Phylogenetic lineages in the Botryosphaeriaceae. Grasso FM, Marini M, Vitale A, Firrao G, Granata G (2012) Canker Studies in Mycology 55|# M # and dieback on Platanus acerifolia caused by Diaporthe scabra. / ]J J @Z “ AE ZZ A X@ V Forest Pathology 42| *!M *# D, et al. !* H ] X V| #_*M#$$ Guarnaccia V, Vitale A, Cirvilleri G, Aiello D, Susca A, et al. (2016) Persoonia: 35|`‡M#_ Characterisation and pathogenicity of fungal species associated /„H/zHZ;]X*$$$X+zE ' % < ]/A' European Journal of Plant Pathology 146|$`#M$_`

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