Three New Species of Corynespora from Indonesia

Abstract

Corynespora hamata, C. acalyphae and C. gracilis are described and illustrated as new species, based on collections made in Indonesia.

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''
Phyllosticta citriasiana sp. nov., the cause of Citrus tan spot of Citrus maxima in
Asia
Wulandari, N.F.1,2,3, To-anun, C1., Hyde, K.D.4,2, Duong, L.M.5, de Gruyter, J.6*, Meffert,
J.P.6, Groenewald, J.Z.7 and Crous, P.W.7
'
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'
["&%#A%,)7' UE!E7' @3L%#"#7' :E7' T/A+7' ME(E7'("3#$7' QE;E7' A+' \,"/.+,7' J E7' ;+44+,.7' JE5E7' \,3+#+]%&A7' JE^E' %#A' :,3"-7'
5E[E'_>??P`E Phyllosticta citriasiana'-1E'#3*E7'.6+'8%"-+'34':).,"-'.%#'-13.'34'Citrus maxima')#'9-)%E'!"#$%&'()*+,-)./'
DNa'>DLDPE'
'
Guignardia'citricarpa7' .6+' 8%"-%&' %$+#.' 34' :).,"-' G&%8K' I13.7' )-' -"Fb+8.' .3' 16/.3-%#).%,/' &+$)-&%.)3#' )#' .6+' V",31+%#'
<#)3#' %#A' .6+' <EIE9E' @6)-' -1+8)+-' )-' 4,+c"+#.&/' 83#4"-+A' ]).6' GE' mangiferae7' ]6)86' )-' %' #3#L1%.63$+#)87' %#A' )-'
83223#&/')-3&%.+A'%-'%#' +#A316/.+'4,32'8).,"-'4,").-'%#A'%'])A+',%#$+'34'3.6+,'63-.-E'B+8+#.&/7'#+8,3.)8'-13.-'-)2)&%,'
.3'.63-+'8%"-+A'F/' GE citricarpa']+,+'3F-+,*+A'3#'4,").'34'Citrus maxima')#.+,8+1.+A')#'83#-)$#2+#.-'+Z13,.+A'4,32'
9-)%E'H#'.6+-+'-13.-7'1/8#)A)%'%#A'83#)A)%'34'%'Guignardia'-1+8)+-'8&3-+&/',+-+2F&)#$'GE citricarpa']+,+'3F-+,*+A7'%#A'
.6+,+43,+'2+%-",+-']+,+' .%K+#' 43,' .6+' 83#-)$#2+#.-')#'&)#+']).6'.6+'V",31+%#'<#)3#'&+$)-&%.)3#'43,'GE citricarpaE'@3'
A+.+,2)#+'.6+')A+#.)./'34'.6+'8%"-%&'3,$%#)-2'3#'.6)-'#+]'63-.7'4"#$%&')-3&%.+-']+,+'-"Fb+8.+A'.3'(U9'-+c"+#8+'%#%&/-)-'
34' .6+' )#.+,#%&' .,%#-8,)F+A' -1%8+,' ,+$)3#' _H@I07' =ECI7' H@I>`7' .,%#-&%.)3#' +&3#$%.)3#' 4%8.3,' 0L%&16%' _@V!0`' %#A' %8.)#'
$+#+-E'9'832F)#+A'16/&3$+#+.)8'.,++',+-3&*+A'.6,++'-1+8)+-'83,,+&%.)#$'.3'GE citricarpa7'GE mangiferae'%#A'%'1,+*)3"-&/'
"#A+-8,)F+A'-1+8)+-7' Phyllosticta citriasiana'-1E'#3*E7'8&3-+&/',+&%.+A'.3'GE citricarpaE';3,163&3$)8%&&/'PE citriasiana'
8%#'F+'A)-.)#$")-6+A'4,32'GE citricarpa'F/'6%*)#$'&%,$+,'83#)A)%7'&3#$+,'83#)A)%&'%11+#A%$+-7'%#A')#'#3.'1,3A"8)#$'%#/'
A)44"-+'/+&&3]'1)$2+#.']6+#'8"&.)*%.+A'3#'3%.2+%&'%$%,'_Y9`E'!",.6+,23,+7').')-'A)-.)#$")-6%F&+'4,32'GE mangiferae'F/'
6%*)#$'-2%&&+,'83#)A)%7']).6'%' #%,,3]+,'2"83)A'-6+%.6E'H#'8"&.",+7'83&3#)+-'34'PE citriasiana'8%#'%&-3'F+'A)-.)#$")-6+A'
4,32' GE citricarpa' %#A' GE mangiferae'F/'F+)#$'A%,K+,'-6%A+-'34'$,+/'%#A' F&%8K' 3#' Y97' 2%&.' +Z.,%8.' %$%,' _;V9`7'
13.%.3LA+Z.,3-+'%$%,7'%#A'83,#2+%&'%$%,E'!",.6+,23,+7'8"&.",+-'34'PE citriasiana'%86)+*+A'31.)2%&'$,3].6'%4.+,'>']++K-'
%.'>0d>Re:7'%#A'8+%-+A'.3'$,3]'%.'D?dDDe:E'H#'83#.,%-.7'83&3#)+-'34'GE citricarpa'%#A'GE mangiferae'%86)+*+A'31.)2%&'
$,3].6' %.' >RdD?e:7' %#A' 8+%-+A' .3' $,3]' %.' D?dDOe:E' :3&3#)+-' 34' PE citriasiana' %&-3' $,+]' 4%-.+,' .6%#' .63-+' 34' GE
citricarpa'%#A'GE mangiferae'3#'Y9'%#A';V9E'Phyllosticta citriasiana' %11+%,-' .3' F+' %' 6%,24"&' 1%.63$+#' 34' Citrus
maxima7'8%"-)#$'%'.%#' -13.'3#'4,").7' "#A+,&)#)#$'.6+' #++A'43,'4",.6+,'-",*+/-'%#A' ,+-+%,86'.3'A+.+,2)#+').-'A)-.,)F".)3#'
%#A'63-.',%#$+E'
'
Key words:':).,"-'G&%8K'I13.7'Guignardia7'c"%,%#.)#+7'23&+8"&%,'16/&3$+#/7'Phyllosticta7'-/-.+2%.)8-E'
'
'
Article Information
B+8+)*+A'0='U3*+2F+,'>??C'
988+1.+A'0='(+8+2F+,'>??C'
5"F&)-6+A'3#&)#+'0'J%#"%,/'>??P'
*:3,,+-13#A)#$'%".63,a'JE'_T%#-`'A+'\,"/.+,f'+L2%)&a'bEA+E$,"/.+,g2)#&#*E#&'
'
Introduction
'
[6+,+' K#3]#7' %#%23,16-' 34' Guignardia'
_Botryosphaeriaceae`' ,+-)A+' )#' Phyllosticta'
_X%#' A+,' 9%7' 0PRDf' 5"#).6%&)#$%27' 0PRNf' X%#'
A+,' 9%' %#A' X%#+*7' >??>`E'Guignardia'-1+8)+-'
6%*+'34.+#' F++#',+83,A+A' %-'+#A316/.+-7' 1&%#.'
1%.63$+#-' %#A' -%1,3F+-' _G%%/+#' et alE7' >??>f'
\&)+#K+LG&%#83' et alE7' >??>f' B3A,)$"+-' et alE7'
>??Nf'T"%#$'et alE7'>??Cf' Ih#86+i' ;h,c"+i' et
alE7' >??Cf' @63#$K%#.6%' et alE7' >??C`E' ;%#/'
Guignardia'-1+8)+-'8%"-+'&+%4'F&3.86'%#A'F&%8K'
-13.-' 3#' 4,").-' 34' *%,)3"-' 1&%#.-' _B%%F+' et alE7'
0PC0f' \&)+#K+LG&%#83' et alE7' >??>`E' !+]'
-."A)+-7' 63]+*+,7' 6%*+' 438"-+A' 3#' .6+' 16/&3L
$+#+.)8' ,+&%.)3#-6)1-' %23#$' Guignardia'%#A'
>D'

Phyllosticta' -1+8)+-7' %#A' "#8+,.%)#./' ,+2%)#-'
1+,.%)#)#$' .3' -1+8)+-' F3"#A%,)+-' %#A' 63-.'
,%#$+-' 34' 23-.' .%Z%' _YK%#+' et alE7' >??0f'
G%%/+#' et alE7' >??>f' :,3"-' et alE7' >??O`E'
I1+8)+-' 34' Guignardia'%,+'83223#&/'K#3]#'
3#&/'4,32'.6+),'%#%23,16-E'@6+')A+#.)4)8%.)3#'34'
Phyllosticta'%#%23,16-')-'6)$6&/'1,3F&+2%.)87'%-'
4+]'86%,%8.+,-'%,+'%*%)&%F&+'.3'-+1%,%.+'A)44+,+#.'
-1+8)+-E' I32+.)2+-' %' -)#$&+' A)44+,+#8+' )#' 83#)L
A)%&' 23,163&3$/7' -"86' %-' .6+' .6)8K#+--' 34' .6+'
2"83)A' &%/+,' -",,3"#A)#$' .6+' 83#)A)"27' 3,' %1L
1+#A%$+' -6%1+' %#A' -)i+7' ,+-"&.+A' )#' %".63,-'
A+-8,)F)#$' A)44+,+#.' -1+8)+-E' X%#' A+,' 9%' %#A'
X%#+*' _>??>`' ,+*)-+A' .6+' >PDO' .%Z%' A+-8,)F+A'
)#' .6+' $+#"-' Phyllosticta7' %88+1.)#$' 0N0' -1+L
8)+-E' ;%#/' 3.6+,' Phyllosticta' -1+8)+-' ]+,+'
832F)#+A' )#.3' $+#+,%' -"86' %-' Ascochyta7'
Coleophoma7' Fusicoccum7' Leptodothiorella7'
Phoma %#A'Phomopsis7'.3'#%2+'F".'%'4+]E''
B+8+#.' -."A)+-' 3#' Guignardia %#A' Phyl-
losticta'6%*+')&&"-.,%.+A'.6+'83#4"-)3#'.6%.'-.)&&'
+Z)-.-'-",,3"#A)#$'-1+8)+-'83#8+1.-')#'&).+,%.",+E'
!3,' )#-.%#8+7' F3.6' Guignardia endophyllicola'
%#A'GE mangiferae'6%*+'F++#'&)#K+A'.3'PE capi-
talensis'%-'%#%23,16'_YK%#+'et alE7'>??0`E' @6)-'
2%..+,' ]%-' ,+-3&*+A' F/' G%%/+#' et alE' _>??>`7'
]63'-"88+--4"&&/' "-+A' H@I'(U9' -+c"+#8+' A%.%'
.3'-63]' .6%.' GE endophyllicola']%-'83#-1+8)4)8'
]).6'GE mangiferaeE'@3'A%.+'.6+'H@I'A32%)#'6%-'
86)+4&/' F++#' "-+A' 43,' -1+8)+-' A)-8,)2)#%.)3#' )#'
.6)-'$,3"1'_V*+,+..'%#A'B++-L\+3,$+7'>??O`7'%#A'
4",.6+,' ]3,K' ,+2%)#-' .3'A+.+,2)#+')4'%'2"&.)L
$+#+' 16/&3$+#+.)8' %11,3%86' ]3"&A' #3.' ,+-3&*+'
23,+'8,/1.)8'.%Z%E'''
:).,"-'G&%8K'I13.7'8%"-+A'F/'Guignardia
citricarpa _%#%23,16'Phyllosticta citricarpa`')-'
,+$"&%.+A'%-' %'c"%,%#.)#+' 1+-.' )#'.6+' V",31+%#'
<#)3#'%#A'.6+'<EIE9E'H.'388",-')#'%'#"2F+,'34'
83"#.,)+-' )#' I3".6+%-.' 9-)%7' 94,)8%7' I3".6'
92+,)8%7'%#A'9"-.,%&)%7'%#A'8%#'F+'A)--+2)#%L
.+A' F/' 2+%#-' 34' )#4+8.+A' 4,").' 3,' *+$+.%.)*+'
1&%#.'2%.+,)%&E'@]3'A)-.)#8.'Guignardia'-1+8)+-'
%,+'%--38)%.+A']).6'8).,"-7'#%2+&/'.6+'1%.63$+#'
GE citricarpa7'%#A'.6+'+#A316/.+'GE mangiferae
_;+/+,' et alE7' >??0f' G%%/+#' etE alE7' >??>f'
V*+,+..'S'B++-L\+3,$+7'>??Of'G%&A%--%,)'et alE7'
>??C`E' G%-+A' 3#' .6+' H@I' -+c"+#8+-7' -+*+,%&'
5:B' A+.+8.)3#' 2+.63A-' 6%*+' F++#' A+*+&31+A'
43,' A+.+8.)3#' 34' GE citricarpa' _G3#%#.-' et alE7'
>??Df' ;+/+,' et alE7' >??Of' 5+,+-' et alE7' >??Rf'
X%#' \+#.L5+&i+,' et alE7' >??R`E' B+8+#.&/7'
#+8,3.)8' -13.-' -)2)&%,' .3' .63-+' 8%"-+A' F/' GE
citricarpa']+,+'3F-+,*+A'3#'4,").'34'Citrus
maxima7' )#.+,8+1.+A' )#' 83#-)$#2+#.-' +Z13,.+A'
4,32'9-)%E'H#'.6+-+'-13.-7'1/8#)A)%'%#A'83#)A)%'
34'%'Guignardia'-1+8)+-7'8&3-+&/',+-+2F&)#$'GE
citricarpa7' ]+,+' 3F-+,*+A' %#A' .6+,+43,+7' 2+%L
-",+-' ]+,+' .%K+#' 43,' .6+' 83#-)$#2+#.-' )#' &)#+'
]).6' .6+' V<' &+$)-&%.)3#' 43,' Guignardia citri-
carpaE''
G/' .+-.)#$' -+*+,%&' 5:B' 2+.63A-' A+*+L
&31+A'43,' A+.+8.)3#'34' GE citricarpa'3#'&+-)3#-'
34' Citrus maxima' %#A' )-3&%.+-' %.' .6+' (".86'
5&%#.' 5,3.+8.)3#' I+,*)8+7' .6+' ,+%&L.)2+' 5:B'
2+.63A' A+*+&31+A' F/' X%#' \+#.L5+&i+,' et alE'
_>??R`' 4%)&+A' .3' 1,3*)A+' %#/' %21&)4)8%.)3#7'
]6)86' ]%-' A"+' .3' -+c"+#8+' A)*+,$+#8+' )#' .6+'
H@I' ,+$)3#E' @6+,+43,+7' %' 83#*+#.)3#%&' 5:B'
%--%/' )-' #3,2%&&/' ,+8322+#A+A' 43,' .6+' A)%$L
#3-)-'34'GE citricarpa'3#'CE maxima'4,").E'@6)-'
]%-' 83#4),2+A' )#' %' 8321%,%.)*+' -."A/' ]).6' %'
83#*+#.)3#%&'5:B'%--%/'A+*+&31+A'F/'G3#%#.-'
et alE' _>??D`' 3#' )-3&%.+-' 3F.%)#+A' 4,32' CE
maxima' _HEBE' T+",#+2%#L*%#' G,3"]+,-6%*+#7'
"#1"F&E'A%.%`E''
@3'A+.+,2)#+'.6+')A+#.)./'34'.6+'Guignar-
dia' -1+8)+-' %--38)%.+A' ]).6' .%#' -13.' 34' Citrus
maxima'4,").')#.+,8+1.+A'4,32'9-)%7'4"#$%&'
)-3&%.+-' ]+,+' -"Fb+8.+A' .3' (U9' -+c"+#8+'
%#%&/-)-' 34' .6+' )#.+,#%&' .,%#-8,)F+A' -1%8+,'
_H@I07' =ECI7' H@I>`' ,+$)3#' %#A' 1%,.)%&' .,%#-&%L
.)3#'+&3#$%.)3#'4%8.3,'0L%&16%'_@V!0`'%#A'%8.)#'
_9:@`' $+#+' -+c"+#8+-E' !",.6+,' %)2-' ]+,+' .3'
)#*+-.)$%.+' .6+' -1+8)+-'F3"#A%,)+-'34'9-)%#'
)-3&%.+-' 8321%,+A' .3' )-3&%.+-' )A+#.)4)+A' %-' GE
citricarpa'%#A'GE'mangiferae7'%#A'.3'A+.+,2)#+'
]6)86' $+#+-' %,+' "-+4"&' )#' A)-.)#$")-6)#$' )-3L
&%.+-'34'Guignardia'%.'.6+'-1+8)+-'&+*+&E'
'
Material and Methods
'
Isolates
:"&.",+-' ]+,+' 3F.%)#+A' 4,32' .6+' :GI'
!"#$%&' G)3A)*+,-)./' :+#.,+' _:GI`7' <.,+86.7'
@6+' U+.6+,&%#A-7' %#A' .6+' 5&%#.' 5,3.+8.)3#'
I+,*)8+' _5(`7' [%$+#)#$+#7' @6+' U+.6+,&%#A-E'
@3' -"11&+2+#.' .6+' A%.%-+.' ]).6' 23,+' -1+8)+-7'
-)#$&+' %-83-13,+' %#A' 83#)A)%&' )-3&%.+-' ]+,+'
,+-1+8.)*+&/' 2%A+' 4,32' Guignardia'%#A'
Phyllosticta'4,").)#$'F3A)+-'388",,)#$'3#'&+%*+-'
%#A' 4,").' 34' A)*+,-+' 63-.' 1&%#.-' _@%F&+' 0`E'
:3&3#)+-']+,+'+-.%F&)-6+A'3#' >' j'2%&.'+Z.,%8.'
%$%,' 1&%.+-' _;V9f' I)$2%L9&A,)86' :6+2)+7'
^])b#A,+86.7' @6+' U+.6+,&%#A-`' F/' "-)#$' .6+'
>N'

!"#$%&'()*+,-)./'
''
.+86#)c"+-' %-' +Z1&%)#+A' )#' :6++]%#$K33#' et
alE'_>??C`E'
'
Molecular phylogeny
(U9' +Z.,%8.)3#' ]%-' A3#+' "-)#$' .6+'
<&.,%:&+%#k' ;)8,3F)%&' (U9' M).' _;Y' G)37'
:%,&-F%A7' :97' <I9`' %883,A)#$' .3' 2%#"4%8L
.",+,l-' 1,3.383&E' @6+' 1,)2+,-' XP\' _A+' T33$'
%#A' \+,,).-' *%#' A+#' V#A+7' 0PPC`' %#A' H@IN'
_[6).+' et alE7' 0PP?`' ]+,+' "-+A' .3' %21&)4/' .6+'
)#.+,#%&' .,%#-8,)F+A' -1%8+,' ,+$)3#' _H@I`' 34' .6+'
#"8&+%,' ,)F3-32%&' BU9' 31+,3#7' )#8&"A)#$' .6+'
Dl' +#A' 34' .6+' 0CI' ,BU97'.6+'4),-.')#.+,#%&'
.,%#-8,)F+A'-1%8+,',+$)3#7'.6+'=ECI',BU9'$+#+f'
.6+' -+83#A' )#.+,#%&' .,%#-8,)F+A' -1%8+,' ,+$)3#'
%#A' .6+' =l' +#A' 34' .6+' >CI' ,BU9' $+#+E' @3'
,+-3&*+' .%Z%' )#' .6+' GE mangiferae'%#A GE'
citricarpa 8321&+Z' .6+' 1,)2+,-' V!0LR>C!' %#A'
V!0LPCOB' _:%,F3#+' %#A' M36#7' 0PPP`' ]+,+'
"-+A' .3' %21&)4/' 1%,.' 34' .6+' .,%#-&%.)3#' +&3#$%L
.)3#' 4%8.3,' 0LĮ' $+#+' _@V!0`' %#A' .6+' 1,)2+,-'
9:@L=0>!' %#A' 9:@LRCDB' _:%,F3#+' %#A'
M36#7' 0PPP`' ]+,+' "-+A' .3' %21&)4/' 1%,.' 34' .6+'
%8.)#' $+#+' _9:@`E' 921&)4)8%.)3#' 83#A).)3#-'
43&&3]+A' 9,i%#&3"' et alE' _>??C`E' 921&)83#-'
]+,+'-+c"+#8+A'"-)#$'F3.6'5:B'1,)2+,-']).6'%'
G)$(/+' @+,2)#%.3,' :/8&+' I+c"+#8)#$' M).' *E'
DE0' _911&)+A' G)3-/-.+2-7' !3-.+,' :)./7' :97'
<I9`'%883,A)#$' .3' .6+'2%#"4%8.",+,l-' )#-.,"8L
.)3#-7'%#A'-+c"+#8+-']+,+'%#%&/-+A'3#'%#'9GH'
5,)-2' DR??' (U9' I+c"+#8+,' _5+,K)#LV&2+,7'
U3,]%&K7'!3-.+,':)./7':97'<I9`E'
I+c"+#8+-' ]+,+' 2%#"%&&/' %&)$#+A' "-)#$'
I+c"+#8+' 9&)$#2+#.' VA).3,' *E' >E?%00' _I+L9&f'
B%2F%".7'>??>`' F/')#-+,.)#$' $%1-E'56/&3$+#+L
.)8'%#%&/-+-' 34' .6+'%&)$#+A' -+c"+#8+'A%.%' ]+,+'
1+,43,2+A' ]).6' 59<5' _56/&3$+#+.)8' 9#%&/-)-'
<-)#$' 5%,-)23#/`' *E' NE?F0?' _I]3443,A7' >??D`'
%-' +Z1&%)#+A' F/' 9,i%#&3"' et alE' _>??R`E' @,++'
&+#$.6' _@Q`7' 83#-)-.+#8/' )#A+Z' _:H`7' ,+.+#.)3#'
)#A+Z'_BH`'%#A',+-8%&+A'83#-)-.+#8'/)#A+Z'_B:`'
]+,+' 8%&8"&%.+A' %#A' .6+' ,+-"&.)#$' .,++-' ]+,+'
1,)#.+A' ]).6' @,++X)+]' *E' 0EOEO' _5%$+' 0PPO`E'
U+]' -+c"+#8+-' ]+,+' &3A$+A' )#' \+#G%#K' %#A'
.6+' %&)$#2+#.' %#A' 16/&3$+#+.)8' .,++' )#@,++L
G9IV' _]]]E.,++F%-+E3,$`E' Botryosphaeria
obtusa _H@I'm'9nPR>0?=7'@V!0'm'(o>C?N0P7'
9:@' m' 9nPR>000`' ]%-' "-+A' %-' 3".$,3"1' )#'
.6+'16/&3$+#+.)8'%#%&/-+-E'
'
Morphology
:"&.",+-' ]+,+' $,3]#' 3#' >j' .%1L]%.+,'
%$%,' -"11&+2+#.+A' ]).6' -.+,)&+' 1)#+' #++A&+-'
_[95`' _:,3"-' et alE7' >??O`7' 43,' 2)8,3-831)8'
+Z%2)#%.)3#E' 5,+1%,%.)3#-' ]+,+' 23"#.+A' )#'
83..3#LF&"+'&%8.316+#3&'3,'8&+%,'&%8.)8'%8)A7'%#A'
-."A)+A'F/'2+%#-'34'%'&)$6.'2)8,3-831+'_u'0???'
2%$#)4)8%.)3#`E'@6+' P='j' 83#4)A+#8+' )#.+,*%&-'
]+,+' A+,)*+A' 4,32' D?' 3F-+,*%.)3#-' 34' -13,+-'
43,2+A' 3#' [957' ]).6' +Z.,+2+-' $)*+#' )#'
1%,+#.6+-+-E'9&&' 8"&.",+-'3F.%)#+A' )#' .6)-' -."A/'
%,+' 2%)#.%)#+A')#' .6+' 8"&.",+' 83&&+8.)3#' 34' .6+'
:GI' _@%F&+' 0`E' :3&3#/' 83&3",-' _-",4%8+' %#A'
,+*+,-+`' ]+,+' %--+--+A'%4.+,' $,3].6' 3#' 43",'
A)44+,+#.' 2+A)%7' 13.%.3LA+Z.,3-+' %$%,' _5(9`7'
83,#2+%&'%$%,'_:;9`7';V9'%#A'3%.2+%&'%$%,'
_Y97'\%2-'et alE7 >??R`'"-)#$'.6+'83&3",'86%,.-'
34'B%/#+,' _0PR?`E'' B%A)%&'8"&.",%&' $,3].6' ,%.+'
]%-'A+.+,2)#+A'%4.+,'>']K'%.'>R'e:')#'.6+'A%,KE'
:%,A)#%&' .+21+,%.",+' ,+c"),+2+#.-' 43,' $,3].6'
]+,+' A+.+,2)#+A' %4.+,' >' ]K' F/' )#8"F%.)#$'
,+1,+-+#.%.)*+'-.,%)#-' _.6,++'-.,%)#-' 1+,' -1+8)+-'
%.' +%86' .+21+,%.",+7' ,+-1+8.)*+&/`' %.' 0>'
A)44+,+#.' .+21+,%.",+-'_4,32'OdDPe:')#'De:'
)#.+,*%&-`' )#' .6+' A%,KE' @6+' #32+#8&%.",%&'
#3*+&./' %#A' A+-8,)1.)3#' ]%-' A+13-).+A' )#'
;/83G%#K'_]]]E;/83G%#KE3,$`E'
'
Results
Phylogeny
@6+' 2%#"%&&/' %Ab"-.+A' 832F)#+A' _H@I7'
@V!0' %#A' 9:@`' %&)$#2+#.' 83#.%)#+A' O?' .%Z%'
_)#8&"A)#$' .6+' 3".$,3"1' -+c"+#8+`' %#A7' 34' .6+'
0?P>' 86%,%8.+,-' "-+A' )#' .6+' 16/&3$+#+.)8'
%#%&/-)-7'>C?' ]+,+'1%,-)23#/L)#43,2%.)*+7'D?0'
]+,+' *%,)%F&+' %#A' 1%,-)23#/L"#)#43,2%.)*+7'
%#A' =00' ]+,+' 83#-.%#.E' U+)$6F3",Lb3)#)#$'
%#%&/-)-'"-)#$'.6+'.6,++'-"F-.).".)3#' 23A+&-' 3#'
.6+' -+c"+#8+' A%.%' /)+&A+A' .,++-' ]).6' )A+#.)8%&'
.313&3$/'%#A'-)2)&%,'F33.-.,%1'*%&"+-E'Y#&/'.6+'
4),-.' 0???' +c"%&&/' 23-.' 1%,-)23#)3"-' .,++-'
]+,+'-%*+A'4,32'.6+'6+",)-.)8'-+%,86'%#A'3#+'34'
.6+-+')-'-63]#')#'!)$E'0'_@Q'm'PN=7':H'm'?ECNR7'
BH'm'?EPR07'B:'m'?EC>>`E'H#A)*)A"%&'$+#+'.,++-'
,+-3&*+A' .6+' -%2+' 8&%A+-' 1,+-+#.+A' )#' !)$E' 0'
%#A'3#&/'A)44+,+A']).6',+$%,A'.3' .6+' 1&%8+2+#.'
34'PE hypoglossi7' PE spinarum7'GE vaccinii'%#A'
Guignardia'-1E'-.,%)#':GI'0???PC'_A%.%'#3.'
-63]#`E' @6,++' ]+&&L-"113,.+A' 8&%A+-' ,+1,+L
-+#.)#$' .6+' 8).,"-' )-3&%.+-' 83"&A' F+' ,+-3&*+AE'
:&%A+' 0 83#-)-.+A' 34' )-3&%.+-' )A+#.)4)+A
>='

Table 1.'(+.%)&-'34'Guignardia'%#A'Phyllosticta')-3&%.+-')#*+-.)$%.+A'A",)#$'.6)-'-."A/E'
'
Species Original
identification
Strain no.1 Substrate Country Collector GenBank no. (ITS, TEF1, ACT)2
Guignardia citricarpa :GI'0?>DN=' Citrus aurantium'
_B".%8+%+`7'&+-)3#-'3#'
1++&
G,%i)&' p' !J=DCD007'!J=DCDOP7'!J=DCN>R'
Guignardia citricarpa :GI'0?>DRDf'
5('PPqP00DCD'
Citrus aurantium'
_B".%8+%+`7'F&%8K'-13.'3#'
4,").
G,%i)&' p' !J=DCD0>7'!J=DCDR?7'!J=DCN>C'
Guignardia citricarpa :GI'0?>DRNf'
5('PPqP00=0P'
Citrus aurantium'
_B".%8+%+`7'F&%8K'-13.'3#'
4,").
G,%i)&' p' !J=DCD0D7'!J=DCDR07'!J=DCN>P'
Guignardia citricarpa :GI'000E>?f'
(I;'D=0N'
p' p' p' !J=DCD0N7'!J=DCDR>7'!J=DCND?'
Guignardia citricarpa :GI'0>?NCPf'
5('
?Nq?0CNNCPR'
Citrus limon'_B".%8+%+` G,%i)&' JE'A+'\,"/.+,' !J=DCD0=7'!J=DCDRD7'!J=DCND0'
Guignardia citricarpa :GI'0>>DCN' Citrus limon'_B".%8+%+` I3".6'94,)8%' ;E'@,".+,' !J=DCD0O7'!J=DCDRN7'!J=DCND>'
Guignardia citricarpa :GI'0>>NC>f'
:5:'0NCNC'
Citrus sinensis
_B".%8+%+`7'&+-)3#-'3#'
4,").
^)2F%F]+' QE'T")-2%#' !J=DCD0R7'!J=DCDR=7'!J=DCNDD'
Guignardia citricarpa :GI'C>CEPR' Citrus aurantium'
_B".%8+%+`7'4,").-'%#A'
&+%*+-
G,%i)&' :E'\&)+#K+' !J=DCD0C7'!J=DCDRO7'!J=DCNDN'
Guignardia mangiferae Guignardia heveae :GI'0?0>>C' Nephelium lappaceum'
_I%1)#A%8+%+`7'
A)-83&3",+A'-1)#.+,-
<I9a'T%]%))' ME9E'U)-6)b)2%' !J=DCD0P7'!J=DCDRR7'!J=DCND='
Guignardia mangiferae GE citricarpa :GI'0??0R=' Citrus'-1E'_B".%8+%+`7'
6+%&.6/'&+%*+-
G,%i)&' :E'\&)+#K+' !J=DCD>?7'!J=DCDRC7'!J=DCNDO'
Guignardia mangiferae GE citricarpa :GI'0??0RO' Citrus'-1E'_B".%8+%+`7'
6+%&.6/'&+%*+-
G,%i)&' :E'\&)+#K+' !J=DCD>07'!J=DCDRP7'!J=DCNDR'
Guignardia mangiferae :GI'00=?NO' Myracrodruon
urundeuva
_9#%8%,A)%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>>7'!J=DCDC?7'!J=DCNDC'
Guignardia mangiferae :GI'00=?NR' Aspidosperma
polyneuron
_9138/#%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>D7'!J=DCDC07'!J=DCNDP'
'
>O'

!"#$%&'()*+,-)./'
''
Table 1 (continued). (+.%)&-'34'Guignardia'%#A'Phyllosticta')-3&%.+-')#*+-.)$%.+A'A",)#$'.6)-'-."A/E'
'
Species Original
identification
Strain no.1 Substrate Country Collector GenBank no. (ITS, TEF1, ACT)2
Guignardia mangiferae :GI'00=?NP' Bowdichia nitida
_!%F%8+%+`7'&+%4'3,'F%,K''
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>N7'!J=DCDC>7'!J=DCNN?'
Guignardia mangiferae :GI'00=?=0' Spondias mombin
_9#%8%,A)%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>=7'!J=DCDCD7'!J=DCNN0'
Guignardia mangiferae :GI'00=?=>' Spondias mombin
_9#%8%,A)%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>O7'!J=DCDCN7'!J=DCNN>'
Guignardia mangiferae :GI'00=?=D' Myracrodruon
urundeuva
_9#%8%,A)%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>R7'!J=DCDC=7'!J=DCNND'
Guignardia mangiferae :GI'00=?=O' Anacardium giganteum
_9#%8%,A)%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>C7'!J=DCDCO7'!J=DCNNN'
Guignardia mangiferae :GI'00=?=R' Anacardium giganteum
_9#%8%,A)%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)c"+-' !J=DCD>P7'!J=DCDCR7'!J=DCNN='
Guignardia mangiferae :GI'00=D0D' Myracrodruon
urundeuva
_9#%8%,A)%8+%+`7'&+%4'3,'
F%,K'
G,%i)&' ME!E'B3A,)$"+-' !J=DCDD?7'!J=DCDCC7'!J=DCNNO'
Guignardia mangiferae :GI'00=DN=' Bowdichia nitida
_!%F%8+%+`7'&+%4'3,'F%,K''
G,%i)&' ME!E'B3A,)$"+-' !J=DCDD07'!J=DCDCP7'!J=DCNNR'
Guignardia mangiferae Guignardia -1E :GI'0>DDRNf'
U![L>>?'
Citrus aurantium'
_B".%8+%+`
@6%)&%#A' UE!E'["&%#A%,)' !J=DCDD>7'!J=DCDP?7'!J=DCNNC'
Guignardia mangiferae Guignardia -1E :GI'0>DN?Nf'
U![L>0P'
Musa paradisiaca
_;"-%8+%+`
@6%)&%#A' UE!E'["&%#A%,)' !J=DCDDD7'!J=DCDP07'!J=DCNNP'
Guignardia mangiferae Guignardia -1E :GI'0>DN?=f'
U![L0=N'
Musa acuminata'
_;"-%8+%+`
@6%)&%#A' UE!E'["&%#A%,)' !J=DCDDN7'!J=DCDP>7'!J=DCN=?'
Guignardia mangiferae GE citricarpa :GI'0RDERRf'
:V:@'>CRN'
Citrus aurantiifolia'
_B".%8+%+`7'4,").
U+]'^+%&%#A' p' !J=DCDD=7'!J=DCDPD7'!J=DCN=0'
Guignardia mangiferae PE capitalensis :GI'>>OERRf'
H!Y'D>P0N'
Paphiopedilum callosum'
_Y,86)A%8+%+`7'&+%4'-13.
\+,2%#/' p' !J=DCDDO7'!J=DCDPN7'!J=DCN=>'
' ' ' '
' ' ' '
>R'

Table 1 (continued). (+.%)&-'34'Guignardia'%#A'Phyllosticta')-3&%.+-')#*+-.)$%.+A'A",)#$'.6)-'-."A/E'
'
Species Original
identification
Strain no.1 Substrate Country Collector GenBank no. (ITS, TEF1, ACT)2
Guignardia mangiferae 9@::'D>R=Rf'
5('
?Nq?0CNNP>O'
Citrus limon _B".%8+%+`7'
&+%4
@%)]%#' JE'A+'\,"/.+,' !J=DCDDR7'!J=DCDP=7'!J=DCN=D'
Guignardia mangiferae :GI'0>?NP?f'
5('
?Nq?0CNNPN>'
Citrus paradisi'
_B".%8+%+`7'4,").
<I97'!&3,)A%' JE'A+'\,"/.+,' !J=DCDDC7'!J=DCDPO7'!J=DCN=N'
Guignardia mangiferae PE musarum :GI'00R00C' Musa acuminata'
_;"-%8+%+`
H#A3#+-)%' HE'G"AA+#6%$+#' !J=DCDDP7'!J=DCDPR7'!J=DCN=='
Guignardia mangiferae GE musae :GI'00PR>?f'
:5:'0D?0D'
Musa -1E'_;"-%8+%+` <I9a'T%]%))' HE'G"AA+#6%$+#' !J=DCDN?7'!J=DCDPC7'!J=DCN=O'
Guignardia mangiferae GE musae :GI'0>DDRDf'
U![L>>0'
Musa paradisiaca
_;"-%8+%+`
@6%)&%#A' UE!E'["&%#A%,)' !J=DCDN07'!J=DCDPP7'!J=DCN=R'
Guignardia mangiferae GE philoprina :GI'D=OE=>f'
9@::'00DOC'
Ilex'-1E _9c")43&)%8+%+` p' p' !J=DCDN>7'!J=DCN??7'!J=DCN=C'
Guignardia mangiferae GE philoprina :GI'DRDE=N' Ilex'-1E _9c")43&)%8+%+` p' p' !J=DCDND7'!J=DCN?07'!J=DCN=P'
Guignardia mangiferae GE sansevieriae :GI'0>?N>Cf'
5('
?Nq?0=NDN?>'
Sansevieria'-1E'
_(,%8%+#%8+%+`
U+.6+,&%#A-' JE'A+'\,"/.+,' !J=DCDNN7'!J=DCN?>7'!J=DCNO?'
Guignardia mangiferae Guignardia capsici :GI'000ODC' Capsicum -1E'
_I3&%#%8+%+`7'4,").
(32)#)8%#'
B+1"F&)8'
\E':%,,3&&' !J=DCDN=7'!J=DCN?D7'!J=DCNO0'
Guignardia mangiferae Guignardia'-1E :;<'0D0' Magnolia liliifera
_;%$#3&)%8+%+`7 &+%4'
+#A316/.+
@6%)&%#A' QE;E'("3#$' !J=DCDNO7'!J=DCN?N7'!J=DCNO>'
Guignardia mangiferae Guignardia'-1E :;<'0DP' Magnolia liliifera
_;%$#3&)%8+%+`7 &+%4'
+#A316/.+
@6%)&%#A' QE;E'("3#$' !J=DCDNR7'!J=DCN?=7'!J=DCNOD'
Guignardia mangiferae Guignardia'-1E :;<'0N>' Magnolia liliifera
_;%$#3&)%8+%+`7 &+%4'
+#A316/.+
@6%)&%#A' QE;E'("3#$' !J=DCDNC7'!J=DCN?O7'!J=DCNON'
Guignardia mangiferae GE vaccinii :GI'00NR=0' Vaccinium'-1E'
_V,)8%8+%+`7'&+%4
U+]'^+%&%#A' @E'!&"6+,' !J=DCDNP7'!J=DCN?R7'!J=DCNO='
Guignardia mangiferae GE philoprina :GI'PDRER?' Hedera helix'
_9,%&)%8+%+`7'&+%4'&)..+,'
H.%&/' [E'\%2-' !J=DCD=?7'!J=DCN?C7'!J=DCNOO'
Guignardia psidii :GI'0??>=?' Psidium guajava'
_;/,.%8+%+`7'4,").-
G,%i)&' :E'\&)+#K+' !J=DCD=07'!J=DCN?P7'!J=DCNOR'
' ' ' '
' ' ' '
>C'

!"#$%&'()*+,-)./'
''
Table 1 (continued). (+.%)&-'34'Guignardia'%#A'Phyllosticta')-3&%.+-')#*+-.)$%.+A'A",)#$'.6)-'-."A/E'
'
Species Original
identification
Strain no.1 Substrate Country Collector GenBank no. (ITS, TEF1, ACT)2
Guignardia -1E :GI'0???PC' Citrus'-1E'_B".%8+%+`7'
6+%&.6/'&+%*+-
G,%i)&' :E'\&)+#K+' !J=DCD=>7'!J=DCN0?7'!J=DCNOC'
Guignardia vaccinii :GI'0>OE>>f'
H!Y'D>P00'
Oxycoccus macrocarpus'
_V,)8%8+%+`
<I9' p' !J=DCD=D7'!J=DCN007'!J=DCNOP'
Phyllosticta citriasiana PE citricarpa :GI'0>?NCCf'
5('
?=q?>NDO?0P'
Citrus maxima'
_B".%8+%+`
@6%)&%#A' JE'A+'\,"/.+,' !J=DCD=N7'!J=DCN0>7'!J=DCNR?'
Phyllosticta citriasiana PE citricarpa :GI'0>DDR?f'
5('
?Cq?NN=DRDO'
Citrus maxima'
_B".%8+%+`
X)+.#%2' JE'A+'\,"/.+,' !J=DCD==7'!J=DCN0D7'!J=DCNR0'
Phyllosticta citriasiana PE citricarpa :GI'0>DDR0f'
5('
?Cq?NN=N0RD'
Citrus maxima'
_B".%8+%+`
X)+.#%2' JE'A+'\,"/.+,' !J=DCD=O7'!J=DCN0N7'!J=DCNR>'
Phyllosticta citriasiana PE citricarpa :GI'0>DDR>f'
5('
?Cq?NN=N0P0'
Citrus maxima'
_B".%8+%+`
X)+.#%2' JE'A+'\,"/.+,' !J=DCD=R7'!J=DCN0=7'!J=DCNRD'
Phyllosticta citriasiana PE citricarpa :GI'0>DDPDf'
5('
?Cq?NN=DR>C'
Citrus maxima'
_B".%8+%+`
X)+.#%2' JE'A+'\,"/.+,' !J=DCD=C7'!J=DCN0O7'!J=DCNRN'
Phyllosticta citriasiana PE citricarpa :GI'0>?N>Rf'
5('
?=q?0O=NCP?'
Citrus maxima
_B".%8+%+`
:6)#%' JE'A+'\,"/.+,' !J=DCD=P7'!J=DCN0R7'!J=DCNR='
Phyllosticta citriasiana PE citricarpa :GI'0>?NCOf'
5('
?=q?0POPR=D'
Citrus maxima
_B".%8+%+`
@6%)&%#A' JE'A+'\,"/.+,' !J=DCDO?7'!J=DCN0C7'!J=DCNRO'
Phyllosticta citriasiana PE citricarpa :GI'0>?NCRf'
5('
?=q?D?C0?=D'
Citrus maxima
_B".%8+%+`
:6)#%' ME'B3-+#A%6&L5+.+,-' !J=DCDO07'!J=DCN0P7'!J=DCNRR'
Phyllosticta citriasiana PE citricarpa :GI'0>?NP0f'
5('
?Oq?D0>=?P='
Citrus maxima
_B".%8+%+`
:6)#%' ME'B3-+#A%6&L5+.+,-' !J=DCDO>7'!J=DCN>?7'!J=DCNRC'
Phyllosticta citriasiana PE citricarpa :GI'0>?NC=f'
5('
?Oq?D0>=00O'
Citrus maxima
_B".%8+%+`
:6)#%' ME'B3-+#A%6&L5+.+,-' !J=DCDOD7'!J=DCN>07'!J=DCNRP'
' ' ' '
' ' ' '
>P'

Table 1(continued). (+.%)&-'34'Guignardia'%#A'Phyllosticta')-3&%.+-')#*+-.)$%.+A'A",)#$'.6)-'-."A/E'
'
Species Original
identification
Strain no.1 Substrate Country Collector GenBank no. (ITS, TEF1, ACT)2
Phyllosticta citriasiana PE citricarpa :GI'0>?N>Of'
5('
?Oq?D0>=0D>'
Citrus maxima'
_B".%8+%+`
:6)#%' ME'B3-+#A%6&L5+.+,-' !J=DCDON7'!J=DCN>>7'!J=DCNC?'
Phyllosticta hypoglossi :GI'0?0ER>f'
H!Y'D>P0O'
Ruscus aculeatus'
_B"-8%8+%+`7'&)*)#$'
&+%*+-
H.%&/' [E'\%2-' !J=DCDO=7'!J=DCN>D7'!J=DCNC0'
Phyllosticta hypoglossi :GI'0OREC=' Ruscus hypoglossum'
_B"-8%8+%+`
H.%&/' [E'\%2-' !J=DCDOO7'!J=DCN>N7'!J=DCNC>'
Phyllosticta hypoglossi :GI'NDNEP>' Ruscus aculeatus'
_B"-8%8+%+`7'A+%A'
8&%A3A+-
H.%&/' [E'\%2-' !J=DCDOR7'!J=DCN>=7'!J=DCNCD'
Phyllosticta owaniana :GI'RROEPRf'
:5:'0??P'
Brabejum stellatifolium'
_5,3.+%8+%+`7'&+%4'-13.
I3".6'94,)8%' 9E'A+#'G,++r+#' !J=DCDOC7'!J=DCN>O7'!J=DCNCN'
09@::a'92+,)8%#'@/1+':"&.",+':3&&+8.)3#7'X),$)#)%7'<EIE9Ef':GIa':+#.,%%&F",+%"'*33,'I86)22+&8"&.",+-7'<.,+86.7'@6+'U+.6+,&%#A-f':;<a';)8,3F)3&3$/'I+8.)3#7':6)%#$';%)'
<#)*+,-)./'_;I:;<`7'(+1%,.2+#.'34'G)3&3$/7'!%8"&./'34'-8)+#8+7':6%#$';%)'<#)*+,-)./7'@6%)&%#Af':5:a':"&.",+'83&&+8.)3#'34'5+A,3':,3"-7'63"-+A'%.':GIf':V:@a':3&+88)3#'
V-1%s3&%'A+':"&.)*3-'@)137'<#)*+,-)./'34'X%&+#8)%7'X%&+#8)%7'I1%)#f'(I;a'(I;^7'(+".-86+'I%22&"#$'*3#';)K,3,,$%#)-2+#'"#A'^+&&K"&.",+#'\2FT7'G,%"#-86]+)$7'\+,2%#/f'
H!Ya'H#-.).".+'!3,'!+,2+#.%.)3#7'Y-%K%7'J%1%#f'U![a':"&.",+'83&&+8.)3#'34'U)&%2'!E'["&%#A%,)f'5(a'5&%#.'5,3.+8.)3#'I+,*)8+7'[%$+#)#$+#7'@6+'U+.6+,&%#A-E'
>H@Ia'H#.+,#%&'.,%#-8,)F+A'-1%8+,-'0'%#A'>'.3$+.6+,']).6'=ECI'#,(U9f'@V!0a'1%,.)%&'.,%#-&%.)3#'+&3#$%.)3#'4%8.3,'0L%&16%'$+#+f'9:@a'1%,.)%&'%8.)#'$+#+E'
'
D?'

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''
%-'GE citricarpa' _:).,"-'G&%8K'I13.`E' Guignar-
dia citricarpa']%-'3,)$)#%&&/'A+-8,)F+A'4,32'
9"-.,%&)%E' 9&.63"$6' .6+' -)#$&+' 9"-.,%&)%#'
)-3&%.+'43"#A')#'.6)-'8&%A+'_:GI'000E>?`'1,3*+A'
.3'F+'-.+,)&+7' ).' -.)&&'1,3A"8+A'.6+'86%,%8.+,)-.)8'
/+&&3]'1)$2+#.']6+#'8"&.)*%.+A'3#' Y9E' :&%A+'
>' ,+1,+-+#.+A' .6+' #+]' -1+8)+-7' A+-8,)F+A' 6+,+'
%-'Phyllosticta citriasiana7'%#A'.6"-' 4%,' K#3]#'
4,32' 8).,"-' 8"&.)*%.+A' )#' :6)#%7' @6%)&%#A' %#A'
X)+.#%2E' :&%A+' D' ,+1,+-+#.+A' )-3&%.+-' )A+#.)L
4)+A'%-'GE mangiferae7')#8&"A)#$'-.,%)#-' 4,32'%'
])A+' 63-.' ,%#$+7' #%2+&/' B".%8+%+' _Citrus'
-11E`7' ;"-%8+%+' _Musa' -1E`7' ;/,.%8+%+'
_Psidium guajava`7' 9#%8%,A)%8+%+' _Mangifera
indica`7' I3&%#%8+%+' _Capsicum annuum`7'
(,%8%+#%8+%+' _Sansevieria'-1E`7'%#A'Y,86)A%L
8+%+' _Y,86)A`E' G%-+A' 3#' .6+-+' A%.%7' -+*+,%&'
-1+8)+-' )#8&"A)#$' GE capsici7 GE capitalensis7
GE endophyllicola7 GE heveae7 GE mangiferae7
GE musae7 GE philoprina7 GE sansevieriae7 %#A
GE vaccinii ]+,+' -63]#' .3' F+&3#$' .3' .6+' GE
mangiferae 8321&+ZE @6+-+' A%.%' -"113,.' .6+'
4)#A)#$-'34'G%%/+#'et alE'_>??>`7'&+%A)#$'.3'.6+'
83#8&"-)3#' .6%.' GE mangiferae')-'%'83-23L
13&).%#'-1+8)+-'.6%.'4,+c"+#.&/'388",-')#'&+-)3#-'
]).6'3.6+,7' 1&%#.' 1%.63$+#)8'-1+8)+-E' 9' 4",.6+,'
8",)3"-' ,+-"&.' ]%-' .6+' 4%8.' .6%.' )-3&%.+' :GI'
0???PC' _4,32' %' Citrus'-1E')#'G,%i)&`'8&"-.+,+A'
-+1%,%.+&/7'%11+%,)#$'23,+'8&3-+&/',+&%.+A'.3'GE
spinarum :GI' PDRER? %#A GE vaccinii :GI'
0>OE>>7' -"$$+-.)#$' .6%.' .6+,+' %,+' /+.' 23,+'
"#,+-3&*+A'-1+8)+-'.6%.'388",'3#'CitrusE''
'
Taxonomy
@6,++' ]+&&LA+4)#+A' -1+8)+-' ]+,+' A+&)L
#+%.+A' 3#' Citrus')#'.6+'1,+-+#.'-."A/E'@6+-+'
)#8&"A+' Guignardia mangiferae'_%#%23,16'
Phyllosticta'capitalensis`7'Guignardia citricar-
pa' _%#%23,16' Phyllosticta citricarpa`7' %#A' %'
Phyllosticta' -1+8)+-' .6%.' )-' 23,163&3$)8%&&/'
A)-.)#8.7' %#A' A3+-' #3.' 83,,+&%.+' ]).6' %#/' 3.6+,'
K#3]#'-1+8)+-'1,+-+#.&/'K#3]#')#'\+#G%#K'3,'
3",' 3]#' (U9' -+c"+#8+' A%.%F%-+-E' !3,' .6+-+'
,+%-3#-'.6)-'-1+8)+-')-'#+]&/'A+-8,)F+A'F+&3]E'
'
Phyllosticta citriasiana ["&%#A%,)7' :,3"- S
\,"/.+,7'sp. nov. !)$E'>E'
;/83G%#Ka';G=?CDCRE'
Teleomorpha'<#K#3]#E'
Spermatial statea'Leptodothiorella -1E'
Etymologya'U%2+A'%4.+,').-'63-.7'Citrus7'%#A'83#.)#+#.'34'
3,)$)#7'9-)%E'
Phyllostictae citricarpae' -)2)&)-7' -+A' 83#)A))-'
2%)3,)F"-7'0?d0O't'=dC'u2E''
Y#' [95E Pycnidia' )22+,-+A' .3'
+,"21+#.7' $&3F3-+7' -"F$&3F3-+' .3' +&&)1-3)A%&E'
VZ"A)#$' -13,+L2%--+-' *%,)+A' 1+,' 8"&.",+' 2+L
A)"27'F+)#$'83&3",&+--'%#A'$&3--/'3#':;9'%#A'
;V97'$,+/'%#A'31%c"+'3#'5(97'%#A'83&3",&+--'
%#A' 31%c"+' 3#' Y9' %#A' [95E' Pycnidia' 0>?d
>N?' t' 0>=d>>=' ȝ2f' 1/8#)A)%&' ]%&&' 83#-)-.)#$'
34'-+*+,%&'&%/+,-7'>=dR?'ȝ2'.6)8Kf'3".+,']%&&'34'
1%&+'F,3]#'.3'F,3]#7'.6)8K+#+A'8+&&-'34'textura
angularis .3' globularisf' )##+,' ]%&&' 83#-)-.)#$'
34' 3#+' .3' .]3' 1%&+' F,3]#' 8+&&' &%/+,-7' .6%.'
F+832+' 6/%&)#+' .3]%,A' )#.+,)3,7' textura
angularisE'Ostiole'-)#$&+7'8+#.,%&7'RdC'ȝ2'])A+7'
D?dD>' ȝ2' A++17' %11+%,)#$' 8/&)#A,)8%&' )#' -+8L
.)3#7'83#-)-.)#$' 34'.6)8K+#+A7'A%,KLF,3]#'8+&&-E'
Conidiophores' -"F8/&)#A,)8%&' .3' %21"&&)43,27'
,+A"8+A' .3' 83#)A)3$+#3"-' 8+&&-' 3,' F,%#86+A'
4,32' %' -"113,.)#$' F%-%&' 8+&&7' Rd>=' u' DdO' u2E'
Conidiogenous cells'.+,2)#%&7'-"F8/&)#A,)8%&'.3'
%21"&&)43,2' 3,' -32+]6%.' A3&))43,27' 6/%&)#+7'
-233.67' 83%.+A' )#' %' .6)#' 2"83)A' &%/+,7'
)#83#-1)8"3"-&/' 1,3&)4+,%.)#$' 3#8+' 3,' .])8+'
1+,8",,+#.&/' #+%,' %1+Z7' Rd0R' t' Dd=' ȝ2E'
Conidia'_0?d`0>d0N_d0O`'t'_=d`OdR_dC`'ȝ27'
-3&).%,/7' 6/%&)#+7' %-+1.%.+7' .6)#L' %#A' -233.6L
]%&&+A7' 83%,-+&/' $".."&%.+7' +&&)1-3)A%&' .3'
3F3*3)A7' .%1+,)#$' .3]%,A' %' #%,,3]&/' .,"#8%.+'
F%-+7'+#8&3-+A')#'%'.6)#'2"8)&%$)#3"-'-6+%.67'0'
ȝ2'.6)8K7'%#A'F+%,)#$'%'6/%&)#+7'2"83)A'%1)8%&'
%11+#A%$+7' Rd0?_d0N`' t' 0d>' ȝ27' -.,%)$6.' .3'
4&+Z)F&+7' "#F,%#86+A7' .%1+,)#$' .3]%,A-' %#'
%8".+&/' ,3"#A+A' .)1E' Spermatia' %.' .)2+-'
43,2)#$')#'83#)A)%&'83#)A)32%.%7'6/%&)#+7'F%8)&L
&)43,2'.3'-32+]6%.'+&&)1-3)A7'Dd='t'0d>'ȝ2E
Specimens examineda' @T9HQ9U(7' 3#' 1++&' 4, ").'
34'Citrus maxima _B".%8+%+`'%-'F&%8K'-13.7'>?'Y8.E'>??=7'
JE'A+'\,"/.+,7' :GI' TL>?0C=7' 63&3./1"-7'8"&.",+'+ZL./1+'
:GI' 0>?NCO' m' 5(' ?=q?0POPR=D`f' :THU97' 3#' 4,").' 34'
Citrus maxima7'>' (+8E' >??=7'ME'B3-+#A%6&L5+.+,-7':GI'
0>?NCR'm'5('?Oq?D0>=?P=`E'
Cultural characteristicsa' :3&3#)+-' 3#'
;V9' 4&%.7' ,+$"&%,7' ]).6' +#.),+' +A$+f' -",4%8+'
&+%A+#L$,+/')#' 8+#.,+7' &%*+#A+,L$,+/'%.'2%,$)#7'
%#A' &+%A+#LF&%8K' "#A+,#+%.6E' Y#' 5(9' 4&%.7'
-1,+%A)#$7']).6'4+%.6+,/'2%,$)#7'4&"44/f'-",4%8+'
A%,K' -&%.+LF&"+7' %#A' 3&)*%8+3"-LF&%8K'
"#A+,#+%.6E' Y#' :;9' 4&%.7' ),,+$"&%,7' ]).6'
&3F+AL+A$+f' -",4%8+' $,++#)-6' F&%8K' )#' 8+#.,+7
D0'

Fig. 1. Y#+'34'0???'+c"%&&/'23-.'1%,-)23#)3"-'.,++-'3F.%)#+A'4,32'%'6+",)-.)8'-+%,86']).6'0??',%#A32'.%Z3#'%AA).)3#-'
34'.6+'832F)#+A'-+c"+#8+'%&)$#2+#.'"-)#$'59<5'*E' NE?F0?E'@6+'-8%&+' F%,'-63]-'0??'86%#$+-7'%#A'F33.-.,%1'-"113,.'
*%&"+-' 6)$6+,' .6%#' R?' j' 4,32' 0???',+1&)8%.+-' %,+' -63]#' %.' .6+' #3A+-E' @6)8K+#+A' &)#+-' )#A)8%.+' .6+' -.,)8.' 83#-+#-"-'
F,%#86+-'%#A'.6+'.,++']%-',33.+A'.3'Botryosphaeria obtusa _H@I'm'9nPR>0?=7'@V!0'm'(o>C?N0P7'9:@'m'9nPR>000`E''
'
100 changes
Botryosphaeria obtusa
Phyllosticta owaniana CBS 776.97
CBS
CBS 102374
CBS 102373
CBS 102345
CBS 828.97
CBS 120489
CBS 122384
CBS 122482
CBS 120487
PD 05/01654890
PD 06/03125132
PD 06/03125116
CBS 120486
PD 05/03081053
CBS 123370
CBS 120488
CBS 123393
CBS 123372
CBS 123371
Phyllosticta spinarum CBS 937.70
Guignardia sp. CBS 100098
Guignardia vaccinii CBS 126.22
CBS 167.85
CBS 101.72
CBS 434.92
Guignardia psidii CBS 100250
Guignardia sp. CMU 142
Guignardia philoprina CBS 356.52
Guignardia heveae CBS 101228
Guignardia philoprina CBS 373.54
Guignardia sp. CBS 123405
Guignardia sp. CBS 123404
Guignardia musae CBS 119720
Guignardia mangiferae CBS
Guignardia mangiferae CBS 115053
Guignardia mangiferae CBS 115052
Guignardia mangiferae CBS 115056
Guignardia mangiferae CBS 115313
Phyllosticta musarum CBS 117118
Guignardia vaccinii CBS 114751
Guignardia sansevieriae PD 04/01543402
Guignardia mangiferae CBS 115345
Guignardia capsici CBS 111638
Guignardia mangiferae CBS
Guignardia mangiferae CBS 115051
Guignardia mangiferae CBS 115057
Guignardia mangiferae CBS 115046
Guignardia mangiferae CBS 115049
Phyllosticta capitalensis CBS 226.77
Guignardia mangiferae CBS 115047
Guignardia mangiferae CBS
Guignardia sp. CMU 131
Guignardia sp. CMU 139
Guignardia mangiferae PD 04/01844942
Guignardia mangiferae PD 01844926
Guignardia musae CBS 123373
93
Guignardia citricarpa
Guignardia sp. CBS 123374
Phyllosticta citriasiana
Phyllosticta hypoglossi
G. mangiferae
species complex
100
100
100
72
100
72
89
93
100
100
100
D>'

!"#$%&'()*+,-)./'
''
'
Fig. 2.'Phyllosticta citriasianaE' %d8E' I/21.32-'3#'4,").'34'Citrus maximaE' AE' :3&3#/' 3#'2%&.'+Z.,%8.'%$%,E'+E' 5/8#)A)%'
-13,"&%.)#$' 3#' -.+,)&+' 1)#+' #++A&+-' 3#' .%1L]%.+,' %$%,E' 4d6E' :3#)A)3$+#3"-' 8+&&-' $)*)#$' ,)-+' .3' -3&).%,/' 83#)A)%E' )7' bE'
:3#)A)%']).6'2"83)A'-6+%.6'%#A'%1)8%&'2"8)&%$)#3"-'%11+#A%$+'*)-)F&+E'I8%&+'F%,-'m'0?'u2E'
'
1%&+' 3&)*%8+3"-L$,+/' %.' 2%,$)#7' %#A' &+%A+#L
F&%8K' "#A+,#+%.6E' Y#' Y9' 4&%.7' ),,+$"&%,7' ]).6'
+#.),+' .3' 4+%.6+,/' 2%,$)#7' ]33&/f' -",4%8+'
&+%A+#LF&%8K' )#' 8+#.,+7' 3&)*%8+3"-LF&%8K' %.'
2%,$)#7' %#A' &+%A+#LF&%8K' .3' &+%A+#L$,+/'
"#A+,#+%.6E''
Cardinal temperaturesa'94.+,'>']K')#'.6+'
A%,K'.6+'31.)2"2'$,3].6',%.+' ]%-' 3F-+,*+A' %.'
D?e:' 3#' ;V97' :;9' %#A' Y9' _>>' 22`f' 3#'
5(9'.6)-'388",,+A'%.'>Re:'_ND'22`E';)#)2"2'
$,3].6',%.+' ]%-' 3F-+,*+A'%.' 0=e:'3#';V9' _='
22`7'5(9'_0='22`7':;9'_='22`'%#A'Y9'_O'
22`E' ;%Z)2"2' $,3].6' ,%.+' ]%-' %.' DDe:' 3#'
;V97' :;9' %#A' Y9' _0R' 22`f' 3#' 5(9' .6)-'
388",,+A'%.'DOe:'_DE='22`E''
Notesa' Phyllosticta citriasiana'A)44+,-'
4,32'.6+' .]3' 3.6+,'-1+8)+-' 388",,)#$'3#' 8).,"-'
)#' ).-' 83#)A)%&' A)2+#-)3#-7' 8"&.",+' 86%,%8.+,)-L
DD'

.)8-' %#A' 8%,A)#%&' .+21+,%.",+' $,3].6' ,+c"),+L
2+#.-E' @6)-' -1+8)+-' 6%-' &%,$+,' 83#)A)%' ]6+#'
8321%,+A' .3' Guignardia citricarpa7' %#A' .6"-'
4%,' )-' 3#&/' K#3]#' 4,32' ).-' Phyllosticta' -.%.+E'
@6+' 83#)A)%&' -6+%.6' )-' )#.+,2+A)%.+' F+.]++#'
.6%.' 34' GE citricarpa'%#A'GE mangiferaeE' @6+'
-6+%.6' ).-+&4' )-' ,%.6+,' .6)#7' F+)#$' 23,+' -)2)&%,'
.3' GE citricarpa' .6%#' GE mangiferae7' ]6+,+%-'
.6+'%1)8%&'%11+#A%$+')-'%$%)#'&3#$+,'.6%#')#' GE
citricarpaE' H#' 8"&.",+' 83&3#)+-' %,+' A%,K+,' .6%#'
.6%.' 34' .6+' 3.6+,' .]3' -1+8)+-7' F+)#$' -6%A+-' 34'
&+%A+#L$,+/'.3'&+%A+#LF&%8K')#'%&&'2+A)%'.+-.+AE'
@6+'2%Z)2"2'.+21+,%.",+'43,'$,3].6'388",,+A'
%.' D?dDDe:7' ]6+,+%-' 43,' .6+' 3.6+,' -1+8)+-' .6)-'
]%-' %.' D?dDOe:E' Q%-.&/7' PE citriasiana' 8%#' F+'
A)-.)#$")-6+A' 4,32' GE citricarpa'F/'#3.'1,3L
A"8)#$' %' A)44"-+' /+&&3]' 1)$2+#.' 3#' Y9E'
56/&3$+#+.)8%&&/7' PE citriasiana'8%#'+%-)&/'F+'
A)-.)#$")-6+A' 4,32' GE citricarpa'%#A'GE
mangiferae' F%-+A' 3#' %&&' .6,++' $+#+' ,+$)3#-'
-+c"+#8+AE' G+.]++#' PE citriasiana'%#A'GE
citricarpa7' 0>' 4)Z+A' #"8&+3.)A+' 86%#$+-' %#A' 0'
)#A+&' ]+,+' 3F-+,*+A' 3*+,' O?>' #"8&+3.)A+-'
_)A+#.)./' 34' PRECN' j`' 43,' H@If' ]6+,+%-' @V!0'
83#.%)#+A' R' 4)Z+A' #"8&+3.)A+' 86%#$+-' %#A' >'
)#A+&-' 3*+,' >R0' #"8&+3.)A+-' _)A+#.)./' 34' POEOC'
j`' %#A' 9:@' 6%A' 3#&/' >' 4)Z+A' #"8&+3.)A+'
86%#$+-'3*+,'>=R'#"8&+3.)A+-'_)A+#.)./'34'PPE>>'
j`'_@%F&+'>`E'
Disease symptoms on fruits
!,").' -/21.32-' %,+' -)2)&%,' .3' .63-+'
1,3A"8+A'F/' GE citricarpa7'.6+' 8%"-%&'%$+#.'34'
:).,"-' G&%8K' I13.E' ' @6+/' 2%)#&/' 83#-)-.' 34'
-6%&&3]'&+-)3#-']).6'%'-2%&&'8+#.,%&'$,+/'.3'.%#'
8,%.+,'"-"%&&/']).6'%'A%,K'F,3]#',)27'Dd0?'22'
A)%2E' @6)-' -/21.32' "-"%&&/' %11+%,-' %4.+,' .6+'
4,").'6%-'-.%,.+A'.3',)1+#E'Y4.+#7'F".'#3.'%&]%/-7'
1/8#)A)%' 8%#' F+' -++#' )#-)A+' .6+' -13.-' %-' .)#/'
%#A' -&)$6.&/' +&+*%.+A' F&%8K' A3.-' )#' .6+' $,+/' .3'
.%#' 4)+&AE' 9' 2%$#)4/)#$' $&%--' 3,' A)--+8.)#$'
2)8,3-831+')-'#++A+A'.3'-++'.6+-+'8&+%,&/E'
9#3.6+,'-/21.32' .6%.' 8%#' -32+.)2+-' F+'
3F-+,*+A' %4.+,' 6%,*+-.7' 83#-)-.-' 34' -2%&&' _0dD'
22' A)%2`7' -&)$6.&/' A+1,+--+A' -13.-E' @6+-+'
-13.-' 2%/' F+' $,+/' .3' .%#7' 3,' ,+AA)-67' 3,'
F,3]#)-67'3,' #3.' A)-83&3",+A'%.' %&&E' Y4.+#'.6+/'
6%*+' %' A%,K' ,+A' 3,' F,3]#' ,)2E' 5/8#)A)%' %,+'
3#&/')#8)A+#.%&&/'1,+-+#.')#'.6+-+'&+-)3#-E';%#/'
)#.+,2+A)%.+-' 388",' F+.]++#' .6+-+' -13.-' %#A'
.6+'1,+*)3"-'./1+E'
Discussion
Guignardia citricarpa'%#A'GE mangiferae'
%,+' .]3' ]+&&L+-.%F&)-6+A' -1+8)+-E' Guignardia
citricarpa')-'83#4)#+A' .3'Citrus'-1+8)+-7' %#A')-'
34' )213,.%#8+' )#' *)+]' 34' 16/.3-%#).%,/'
,+c"),+2+#.-' _\&)+#K+LG&%#K3' et alE7' >??>`E'
Guignardia mangiferae' 6%-' F++#' ,+83,A+A' 3#'
2%#/' 63-.-' %#A' )-' %' 83223#' +#A316/.+' 34'
A)*+,-+' ]33A/' 63-.' 1&%#.-' _G%%/+#' et alE7'
>??>`E' @6+,+' 6%-7' 63]+*+,7' F++#' 83#-)A+,%F&+'
83#4"-)3#' %F3".' .6+' )A+#.)4)8%.)3#' 34' .6+-+'
-1+8)+-E';3,163&3$)8%&&/'.6+-+'.]3'-1+8)+-'%,+'
A)-.)#8.E'Guignardia citricarpa'A)44+,-'4,32'GE
mangiferae')#'%-83-13,+'-)i+7'%#%23,16'86%L
,%8.+,-' %#A' 1%.63$+#)8)./E' 9-83-13,+-' 34' GE
citricarpa'_Cd0R't'DE=dC'ȝ2`'%,+'"-"%&&/'&%,$+,'
.6%#'.63-+'34'GE mangiferae'_0?d0>'t'Nd='ȝ2`E'
:3#)A)%' 34' PE' citricarpa'_Pd0?'t'OdR'ȝ2` ' %,+'
&%,$+,'.6%#'.63-+'34'PE capitalensis'_Cd0?'t'Nd='
ȝ2`' _%#%23,16' 34' GE mangiferae`7' %#A' %&-3'
6%*+' %' .6)##+,' -6+%.6E' T3]+*+,' 2)-)A+#.)L
4)8%.)3#' 34' .6+' .]3' 4"#$)'6%-'34.+#'388",,+A'
_V*+,+..'%#A'B++-L\+3,$+7'>??O`E'
H#']6%.'6%-' 1,3*+#' .3'F+'%' 1)*3.%&' 1%1+,'
3#'Guignardia'.%Z3#32/7'G%%/+#'et alE'_>??>`'
"-+A' H@I' -+c"+#8+-' .3' %#%&/-+' Guignardia'
)-3&%.+A' 4,32' Citrus' -11E' 4,32' *%,)3"-' &38%L
.)3#-' .3' )#*+-.)$%.+' .6+' A)-.)#8.)3#' F+.]++#'
1%.63$+#)8'GE citricarpa'%#A'#3#1%.63$+#)8'GE
mangiferaeE'@6+/'A)*)A+A'.6+')-3&%.+-' )#.3' .]3'
A)44+,+#.' $,3"1-' 34' Guignardia'F%-+A'3#'
23,163&3$/' %#A' H@I' -+c"+#8+' A%.%E' @6+' 4),-.'
$,3"1' 8321,)-+A' -.,%)#-' )-3&%.+A' 4,32' F&%8K'
-13.-' %#A' .6+' -+83#A' $,3"1' 8321,)-+A' -.,%)#-'
4,32' Citrus' -11E' %#A' 0C' 3.6+,' 63-.-E' H@I'
%#%&/-+-' 34' .6+' -.,%)#-' 4,32' .6+-+' .]3' $,3"1-'
)#8&"A)#$' ,+4+,+#8+' Guignardia'-+c"+#8+-'
,+-"&.+A' )#' -.,%)#-' 4,32' .6+' 4),-.' $,3"1' F+)#$'
83#-)A+,+A' .3' F+' Guignardia citricarpa'sensu
strictof'.6+'-.,%)#-'4,32'.6+'-+83#A'$,3"1'F+)#$'
83#-1+8)4)8']).6' Phyllosticta capitalensisE' @6+'
-.,%)#-'4,32'.6+'0C'63-.-'3.6+,'.6%#'Citrus']+,+'
$+#+,%&&/' )-3&%.+A' 4,32' 6+%&.6/' &+%*+-7' ]6+,+'
.6+' 4"#$"-' ]%-' 1,+-+#.' %-' %#' +#A316/.+7' %#A'
83)#8)A+#.%&&/'4,32'-13..+A'&+%*+-E'U3#+'34'.6+'
)-3&%.+-' 34' .6+' -+83#A' $,3"1' 8%2+' 4,32' 4,").-'
]).6'8&%--)8%&'F&%8K'-13.-E'@6+/'83#8&"A+A'.6%.
GE mangiferae')-'1,+-+#.'%-'+#A316/.+')#'2%#/'
A)44+,+#.' 63-.-' 34' *%,)3"-' 1&%#.' 4%2)&)+-E' @6+'
23,163&3$)8%&' A)-.)#8.)3#' F+.]++#' .6+
DN'

!"#$%&'()*+,-)./'
''
D='
Table 2.' U"8&+3.)A+' A)44+,+#8+-' %#A' .6+),' F%-+' 13-).)3#-' 3F-+,*+A' )#' .6,++' &38)' F+.]++#' Guignardia citricarpa'%#A'Phyllosticta citriasianaE'
I+c"+#8+-'34' Guignardia citricarpa'-.,%)#':GI'000E>?']+,+'"-+A' %-',+4+,+#8+-'.3'8%&8"&%.+'F%-+' 13-).)3#-7']6)86'A3'#3.')#8&"A+'-1%8+-'8%"-+A'
F/'%&)$#2+#.'$%1-E'U"8&+3.)A+-')#'F3&A'1,)#.'%,+')A+#.)8%&'.3'.6+',+4+,+#8+'-+c"+#8+'%#A'F%-+-')#',3"#A'1%,+#.6+-+-']+,+'#3.'83#-)A+,+A'%-'4)Z+A'
#"8&+3.)A+'86%#$+-'-1+8)4)8'.3'%'-1+8)+-E'I++'@%F&+'0'43,'.6+'A+4)#).)3#'34'.6+'-.,%)#'%#A'&38"-'%FF,+*)%.)3#-'%#A'43,'8321&+.+'-.,%)#')#43,2%.)3#E'
'
ITS1 ITS2 Species'Strain'
68a 77/78c 83b 98a 129a 172a 187a 189b 191a 234b 245a 275a 554a
Guignardia citricarpa :GI'000E>?' G - T G T C C C A C C A G
Guignardia citricarpa :GI'C>CEPR' G - T G T C C C A C C A G
Guignardia citricarpa :GI'0?>DN=' G - T G T C C C A C C A G
Guignardia citricarpa :GI'0?>DRD' G - T G T C C C A C C A G
Guignardia citricarpa :GI'0?>DRN' G - T G T C C C A C C A G
Guignardia citricarpa :GI'0>?NCP' G - T G T C C C A C C A G
Guignardia citricarpa :GI'0>>DCN' G - T G T C C C A C C A G
Guignardia citricarpa :GI'0>>NC>' G - T G T C C C A C C A G
Phyllosticta citriasiana :GI'0>?NCO' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana :GI'0>?NCR' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana :GI'0>?NCC' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana :GI'0>DDR?' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana :GI'0>DDR0' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana :GI'0>DDR>' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana :GI'0>DDPD' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana 5('?=q?0O=NCP?' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana 5('?=q?D?C0?=D' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana 5('?Oq?D0>=00O' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
Phyllosticta citriasiana 5('?Oq?D0>=0D>' 9' \' \' 9' :' @' @' 9' \' 9' @' \' 9'
%'@,%#-).)3#E'
F'@,%#-*+,-)3#E'
8'H#-+,.)3#'q'A"1&)8%.)3#'34'&+%A)#$'#"8&+3.)A+E'
'

Table 2 (continued).' U"8&+3.)A+' A)44+,+#8+-' %#A' .6+),' F%-+' 13-).)3#-' 3F-+,*+A' )#' .6,++' &38)' F+.]++#' Guignardia citricarpa'%#A'Phyllosticta
citriasianaE'I+c"+#8+-'34'Guignardia citricarpa'-.,%)#':GI'000E>?']+,+'"-+A'%-',+4+,+#8+-'.3'8%&8"&%.+' F%-+' 13-).)3#-7' ]6)86'A3'#3.')#8&"A+'-1%8+-'
8%"-+A'F/'%&)$#2+#.'$%1-E'U"8&+3.)A+-')#'F3&A'1,)#.'%,+')A+#.)8%&'.3'.6+',+4+,+#8+'-+c"+#8+'%#A'F%-+-')#',3"#A'1%,+#.6+-+-']+,+'#3.'83#-)A+,+A'%-'4)Z+A'
#"8&+3.)A+'86%#$+-'-1+8)4)8'.3'%'-1+8)+-E'I++'@%F&+'0'43,'.6+'A+4)#).)3#'34'.6+'-.,%)#'%#A'&38"-'%FF,+*)%.)3#-'%#A'43,'8321&+.+'-.,%)#')#43,2%.)3#E'
'
TEF1 Species'Strain'
32b (49)b 74b 80c 107a 129a 150a 173c 218a 228a (263)a (270)a
Guignardia citricarpa :GI'000E>?' A A T T C A T T A T T C
Guignardia citricarpa :GI'C>CEPR' A A T T C A T T A T T @'
Guignardia citricarpa :GI'0?>DN=' A A T T C A T T A T T @'
Guignardia citricarpa :GI'0?>DRD' A A T T C A T T A T T @'
Guignardia citricarpa :GI'0?>DRN' A A T T C A T T A T T @'
Guignardia citricarpa :GI'0>?NCP' A A T T C A T T A T :' @'
Guignardia citricarpa :GI'0>>DCN' A A T T C A T T A T :' @'
Guignardia citricarpa :GI'0>>NC>' A A T T C A T T A T :' @'
Phyllosticta citriasiana :GI'0>?NCO' @' A 9' L' @' \' :' L' \' :' T C
Phyllosticta citriasiana :GI'0>?NCR' @' A 9' L' @' \' :' L' \' :' T @'
Phyllosticta citriasiana :GI'0>?NCC' @' A 9' L' @' \' :' L' \' :' :' @'
Phyllosticta citriasiana :GI'0>DDR?' @' A 9' L' @' \' :' L' \' :' :' @'
Phyllosticta citriasiana :GI'0>DDR0' @' A 9' L' @' \' :' L' \' :' :' @'
Phyllosticta citriasiana :GI'0>DDR>' @' A 9' L' @' \' :' L' \' :' :' @'
Phyllosticta citriasiana :GI'0>DDPD' @' A 9' L' @' \' :' L' \' :' :' @'
Phyllosticta citriasiana 5('?=q?0O=NCP?' @' @' 9' L' @' \' :' L' \' :' T C
Phyllosticta citriasiana 5('?=q?D?C0?=D' @' @' 9' L' @' \' :' L' \' :' T C
Phyllosticta citriasiana 5('?Oq?D0>=00O' @' A 9' L' @' \' :' L' \' :' T C
Phyllosticta citriasiana 5('?Oq?D0>=0D>' @' A 9' L' @' \' :' L' \' :' T C
%'@,%#-).)3#E'
F'@,%#-*+,-)3#E'
8'H#-+,.)3#'q'A"1&)8%.)3#'34'&+%A)#$'#"8&+3.)A+E'
DO'
'
'

!"#$%&'()*+,-)./'
''
DR'
Table 2 (continued).'U"8&+3.)A+'A)44+,+#8+-'%#A'.6+),'F%-+'13-).)3#-'3F-+,*+A')#'.6,++'&38)'F+.]++#'
\")$#%,A)%' 8).,)8%,1%' %#A' 56/&&3-.)8.%' 8).,)%-)%#%E' I+c"+#8+-' 34' \")$#%,A)%' 8).,)8%,1%' -.,%)#' :GI'
000E>?']+,+'"-+A' %-' ,+4+,+#8+-'.3'8%&8"&%.+'F%-+'13-).)3#-7']6)86'A3'#3.')#8&"A+'-1%8+-'8%"-+A'F/'
%&)$#2+#.'$%1-E'U"8&+3.)A+-')#'F3&A'1,)#.'%,+')A+#.)8%&'.3'.6+',+4+,+#8+'-+c"+#8+'%#A'F%-+-')#',3"#A'
1%,+#.6+-+-']+,+'#3.'83#-)A+,+A'%-'4)Z+A'#"8&+3.)A+'86%#$+-' -1+8)4)8' .3' %' -1+8)+-E' I++'@%F&+'0'43,'
.6+'A+4)#).)3#'34'.6+'-.,%)#'%#A'&38"-'%FF,+*)%.)3#-'%#A'43,'8321&+.+'-.,%)#')#43,2%.)3#E'
'
ACT Species'Strain'
(11)a (55)b (101)b 118b (190)a 204a (209)b
Guignardia citricarpa :GI'000E>?' C C T G G T G
Guignardia citricarpa :GI'C>CEPR' C C T G G T G
Guignardia citricarpa :GI'0?>DN=' C C T G G T G
Guignardia citricarpa :GI'0?>DRD' C C T G G T G
Guignardia citricarpa :GI'0?>DRN' @' C T G G T G
Guignardia citricarpa :GI'0>?NCP' C C T G G T G
Guignardia citricarpa :GI'0>>DCN' C C T G G T G
Guignardia citricarpa :GI'0>>NC>' C C T G G T G
Phyllosticta citriasiana :GI'0>?NCO' C C T @' 9' :' G
Phyllosticta citriasiana :GI'0>?NCR' C C T @' G :' G
Phyllosticta citriasiana :GI'0>?NCC' C 9' \' @' 9' :' :'
Phyllosticta citriasiana :GI'0>DDR?' C C T @' 9' :' G
Phyllosticta citriasiana :GI'0>DDR0' C C T @' 9' :' G
Phyllosticta citriasiana :GI'0>DDR>' C C T @' 9' :' G
Phyllosticta citriasiana :GI'0>DDPD' C C T @' 9' :' G
Phyllosticta citriasiana 5('?=q?0O=NCP?' C C T @' G :' G
Phyllosticta citriasiana 5('?=q?D?C0?=D' C C T @' 9' :' G
Phyllosticta citriasiana 5('?Oq?D0>=00O' C C T @' G :' G
Phyllosticta citriasiana 5('?Oq?D0>=0D>' C C T @' G :' G
%'@,%#-).)3#E'
F'@,%#-*+,-)3#E'
8'H#-+,.)3#'q'A"1&)8%.)3#'34'&+%A)#$'#"8&+3.)A+E'
'
%#%23,16-'34'GE citricarpa'%#A'GE mangiferae'
)-' 43"#A' )#' .6+' .6)8K#+--' 34' .6+' 83#)A)%&'
2"8)&%$)#3"-' -6+%.6E' H#' *).,3' GE citricarpa'
-.,%)#-' 1,3A"8+' %' A)-.)#8.' /+&&3]' 1)$2+#.' 3#'
Y9E' T3]+*+,7' 23&+8"&%,' .33&-' %,+' ,+c"),+A' .3'
,%1)A&/')A+#.)4/'.6+-+'.]3'-1+8)+-E'@6+')-3&%.+-'
3F.%)#+A'4,32'Citrus maxima'8&+%,&/',+1,+-+#.'
%' A)44+,+#.' .%Z3#7' 43,' ]6)86' .6+' #%2+' PE
citriasiana' ]%-' )#.,3A"8+AE' Phyllosticta
citrisiana'8%#'F+'A)-.)#$")-6+A'4,32'GE
mangiferae' F/' 6%*)#$' -2%&&+,' 83#)A)%7' ]).6' %'
#%,,3]+,' 2"83)A' -6+%.6E' !",.6+,23,+7' ).' )-'
A)-.)#$")-6%F&+' 4,32' GE citricarpa'F/'6%*)#$'
&%,$+,'83#)A)%7'&3#$+,'83#)A)%&'%11+#A%$+-7'%#A'
#3.' 1,3A"8)#$' %#/' A)44"-+' /+&&3]' 1)$2+#.'
]6+#'8"&.)*%.+A'3#'Y9E'H#' 8"&.",+7' 83&3#)+-' 34'
PE citriasiana'%,+'%&-3'A%,K+,'-6%A+-'34'$,+/'
%#A'F&%8K'3#' Y97' ;V97'5(97'%#A':;9' .6%#'
3F-+,*+A')#'.6+'3.6+,'.]3'-1+8)+-E''
H#'.6+'1,+-+#.'-."A/7'.6,++'-.,%)#-')-3&%.+A'
4,32' Citrus'-11E'_:GI'0??0R=7':GI'0??0RO'
%#A' :GI' 0RDERR`' %&.63"$6' .6+-+' -13,"&%.)#$'
133,&/' 3#' Y97' 1,3A"8+' /+&&3]' 1)$2+#.-' )#'
8"&.",+7' %#A' %,+' -++#' %-' GE citricarpa'sensu
strictoE'T3]+*+,7')#'.6)-'23&+8"&%,'-."A/' .6+-+'
)-3&%.+-'8&"-.+,+A')#'.6+'GE mangiferae'8321&+Z'
_!)$E' 0`E' @6+' 1,+-+#.' -."A/' ]%-' F%-+A' 3#' .6+'
%*%)&%F&+' 8"&.",+-' %.' .6+' :GI' %#A' .6+' (".86'
5&%#.' 5,3.+8.)3#' I+,*)8+' 8"&.",+' 83&&+8.)3#-7'
%#A' -32+' 4,+-6' )-3&%.+-' 3F.%)#+A' 4,32' 9-)%E'
U"2+,3"-' -1+8)+-' 34' Guignardia %#A' Phyl-
losticta',+c"),+'4",.6+,'-."A/7'%-'4+]'6%*+'F++#'
8321%,+A' .6"-' 4%,' 3#' %' 23&+8"&%,' F%-)-E'
9&.63"$6'1,+&)2)#%,/7'3",'A%.%'-"$$+-.'.6%.'.6+'
H@I' &38"-' )-' )#-"44)8)+#.' 43,' -+1%,%.)#$' %&&'
8,/1.)8' .%Z%' )#' Guignardia'_Phyllosticta`E'
T3]+*+,7' H@I' _PRECN' j' )A+#.)./' F+.]++#' PE
citriasiana'%#A'GE citricarpa`'%#A'@V!0'_POEOC'
j' )A+#.)./' F+.]++#' PE citriasiana'%#A'GE
citricarpa`' $%*+' %' F+..+,' -1+8)+-' ,+-3&".)3#'
F+.]++#' PE citriasiana'%#A'GE citricarpa' .6%#'
9:@'_PPE>>' j')A+#.)./' F+.]++#' PE citriasiana'
%#A' GE citricarpa`' )#' .6)-' -."A/E' ;3,+' .,%#L
-).)3#-'.6%#'.,%#-*+,-)3#-']+,+'3F-+,*+A'_@%F&+'
>`' 43,' H@I' %#A' @V!0' %#A' %#' %&23-.' +c"%&'
4,+c"+#8/'43,'9:@E'U3#+'34'.6+'.6,++'$+#+-')#'
.6)-' -."A/' ,+*+%&+A' -)$#)4)8%#.' *%,)%.)3#' )#' .6+'
GE mangiferae'8321&+Z'_A%.%'#3.'-63]#f'#+%,'

DC'
*+,.)8%&'&)#+' )#' !)$E'0`E'!",.6+,' )#*+-.)$%.)3#' )-'
.6"-' 8%&&+A' 43,7' .3' A+.+,2)#+' )4' 3.6+,' &38)'
-"113,.' .6+' 23,163&3$)8%&' *%,)%.)3#' 3F-+,*+A'
%23#$')-3&%.+-')#'GE mangiferaeE''@6+'%11%,+#.'
-/#3#/2/'34'#"2+,3"-'.%Z%'"#A+,'.6)-'+1).6+.7'
.6+,+43,+' _G%%/+#' et alE7' >??>`7' -63"&A' F+'
%88+1.+A']).6'-32+',+-+,*%.)3#E'
Y",' -."A/' 6%-' ,+*+%&+A'.]3'-1+8)+-'34'
Guignardia'.3'8%"-+'A)-+%-+-'34'CitrusE'
Guignardia citricarpa'8%"-+-':).,"-'G&%8K'
I13.')#'I3".6+%-.'9-)%7'94,)8%7'I3".6' 92+,)8%'
%#A'9"-.,%&)%7']6)&+' PE citriasiana')-'1,+-+#.&/'
K#3]#'3#&/'4,32'9-)%7']6+,+').'8%"-+-'%':).,"-'
@%#'I13.'3#'Citrus'maxima'4,").E'G+8%"-+'.6+,+'
%,+' -+*+,%&' Guignardia' -1+8)+-' ]).6' %' -)2)&%,'
83#)A)%&' 23,163&3$/' .6%.' 8%"-+' A)-+%-+' 3#' %'
,%#$+'34'8"&.)*%.+A'1&%#.-7').')-'"#8&+%,']6+.6+,'
.6+-+' 83"&A' ,+1,+-+#.' +).6+,' GE citricarpa'3,'%'
.+&+323,16' 34' PE citriasianaE' !",.6+,' -",*+/-7'
1%.63$+#)8)./' -."A)+-' %#A' 23&+8"&%,' %#%&/-+-'
%,+' .6"-' ,+c"),+A7' .3' ,+-3&*+' .6+' A)-.,)F".)3#7'
63-.',%#$+'%#A')213,.%#8+'34'.6+-+'.]3'-1+8)+-E'
9' -",*+/' 2%/' %&-3' %#-]+,' .6+' c"+-.)3#'
]6+.6+,'.6+,+')-' %' .+&+323,16'34'PE citriasiana'
388",,)#$' )#' 3,86%,A-' )#' .6+' 9-)%#' %,+%' 34'
3,)$)#E''
'
Acknowledgements
'
[+' %,+' $,%.+4"&' .3' G,%2' A+' T331' 43,' 8,).)8%&'
,+%A)#$' .6+' 2%#"-8,)1.E' [+' K)#A&/' .6%#K' M%,)#'
B3-+#A%6&L5+.+,-' ]63' 3F.%)#+A' 2%#/' -.,%)#-' -."A )+A' )#'
.6)-' 1%1+,' *)%' ,3".)#+' )-3&%.)3#-' %.' .6+' 5&%#.' 5,3.+8.)3#'
I+,*)8+E'
'
References
'
9,i%#&3"7';E7' \,3+#+]%&A7'JE^E7'!"&&+,.3#7'BE9E7' 9F+&#7'
VE:E9E7' :%,&)+,7' JE7' ^%1%.+,7' ;E!E7' G"AA+#6%$+#7'
HE[E7' X)&b3+#7' 9E' %#A' :,3"-7' 5E[E' _>??C`E'
;"&.)1&+' $+#+' $+#+%&3$)+-' %#A' 16+#3./1)8'
86%,%8.+,-' A)44+,+#.)%.+' -+*+,%&' #3*+&' -1+8)+-' 34'
Mycosphaerella'%#A',+&%.+A'%#%23,16-'3#'
F%#%#%E'5+,-33#)%'>?a'0PLDRE'
9,i%#&3"7' 9E7' \,3+#+]%&A7' JE^E7' \%2-7' [E7' G,%"#7' <E7'
I6)#7'TE(E' %#A':,3"-7'5E[E'_>??R`E'56/&3$+#+.)8'
%#A' 23,163.%Z3#32)8' ,+*)-)3#' 34' Ramichlo-
ridium'%#A'%&&)+A'$+#+,%E'Studies in Mycology'=Ca'
=RLPDE'
G%%/+#7' BE5E7' G3#%#.-7' 5EJE;E7' X+,K&+/7' \E7' :%,,3&&7'
\E:E7' 9%7' *%#' A+,' TE9E7' [++,A.7' (+' ;E7'
G,3"]+,-6%*+#7' *%#' HEBE7' I86"..+7' \E:E7'
;%886+,3#)7' J,E' [E7' G&%#837' A+' :E\E' %#A'
9i+*+A37' JEQE' _>??>`E' U3#1%.63$+#)8' )-3&%.+-' 34'
.6+' :).,"-' G&%8K' I13.' 4"#$"-7' Guignardia
citricarpa7' )A+#.)4)+A' %-' %' 83-2313&).%#'
+#A316/.+' 34' ]33A/' 1&%#.-7' GE mangiferae'
_Phyllosticta capitalensis`E' 56/.31%.63&3$/' P>a'
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PROOF
Cryptogamie, Mycologie, 2010, 31 (4): 1-000
© 2010 Adac. Tous droits réservés
Guignardia/Phyllosticta species on banana
Nilam F. WULANDARI1,2, Chaiwat TO-ANUN1, Lei CAI 3,
Kamel A. ABD-ELSALAM4,5,6 & Kevin D. HYDE 4,7
1Department of Plant Pathology Faculty of Agriculture, Chiang Mai University,
Chiang Mai, 51200, Thailand
2Microbiology Division, Research Centre for Biology,
Indonesian Institute of Sciences, Cibinong Science Centre Jl. Raya Jakarta,
Bogor KM 46, Cibinong 16911, Indonesia
nilamwulandari@gmail.com
3Key Laboratory of Systematic Mycology & Lichenology, Institute of Microbiology,
Chinese Academy of Sciences, No. 10, North 4th Ring Road West Beijing 100190,
P.R.China No. 14, Xinxi Road, Shangdi, HaiDian, Beijing, 100085, PR China
4Botany and Microbiology Department, College of Science, King Saud University,
Riyadh, Saudi Arabia
5Abdul Rahman Al-Jeraisy, DNA Research Chair, College of Science,
King Saud University, Riyadh, Saudi Arabia
6Plant Pathology Research Institute, Agricultural Research Centre, 9-Gamma St.,
Giza, Egypt
7School of Sciences, Mae Fah Luang University, 333 M. 1. T. Tasud Muang District,
Chiang Rai 57100, Thailand
Abstract –Guignardia musae is the reported causal agent of freckle disease of banana. The
epithet has, however, been introduced on three separate occasions and only one name is
valid. We therefore investigated this problem. We examined the types of G. musae Racib.,
G. musae F. Stevens and G. musae Syd. & P. Syd. and also made fresh collections from
banana in northern Thailand. Guignardia musae Racib. is the earliest name and takes
precedence over the other two names which are hononyms. G. musae F. Stevens is a
different species and therefore a new name G. stevensii Wulandari & K.D. Hyde is
introduced to accommodate it. The name G. sydowiana Trotter has previously been
introduced to accommodate G. musae Syd. & P Syd.; type material is, however,
depauperate. Guignardia musicola Wulandari, L. Cai & K.D. Hyde sp. nov. is introduced as
a new species from Thailand. The three species from banana are compared and their
differences described.
Banana freckle disease / New species / Taxonomy
000_000_Wulandari.fm Page 1 Lundi, 20. décembre 2010 4:09 16

PROOF
2N. F. Wulandari, C. To-Anun, L. Cai, K. A. Abd-Elsalam & K. D. Hyde
INTRODUCTION
Freckle disease occurs on several species and varieties in Musaceae
(Jones & Alcorn, 1982; Jones, 1984, 1993, 1994a, b, 1999; Pitakpaivan, 1985;
Shivas et al., 1996). The causal agent induces freckling on the leaves and fruits,
causing a series of black, raised spots with a sand paper-like texture; this is due
to the protruding pycnidia and/or ascomata. Leaves turn yellow with time and
eventually scenesce. The causal agent of banana freckle is reported to be
Guignardia musae (Aa, 1973; Aa & Vanev, 2002; CABI, 1990, 2005; Chuang,
1981; Dingley et al., 1981; Hwang et al., 1984; Jones & Alcorn, 1982; Jones, 1984,
1993, 1994a, b, 1999; Meredith, 1968; Pitakpaivan, 1985; Ploetz et al., 2003;
Sivanesan, 1984; Shivas et al., 1996; Tsai et al., 1993; Zhou & Xie 1992) and its
anamorph is reported to be Phyllosticta musarum (Aa, 1973; Aa & Vanev, 2002;
Sivanesan, 1984).
The name Guignardia musae has been introduced on three occasions. It
was first introduced by Raciborski (1909) for a fungus on Musa paradisiaca from
Indonesia. This was followed by G. musae F. Stevens from Musa sp. in Hawaii
(Stevens, 1925) and G. musae Syd. & P. Syd from Musa sp. in the Democratic
Republic of Congo (Sydow & Sydow, 1912). The latter two names are homonyms
and thus invalid. In the literature and generally on the world-wide web, the
cause of freckle is listed as Guignardia-Phyllosticta sp. (http://www.pestnet.org/
Summaries/Crops/Plantationcrops/Banana/Fungi/Frecklediseaseofbanana/
tabid/1350/Default.aspx; http://www2.dpi.qld.gov.au/horticulture/7926.html and
http://www.indexfung-orum.org) and the exact name of the species is not often
listed.
Banana freckle occurs worldwide (Table 1). It is common in Asia where
the causative agent is usually listed as Guignardia musae Racib. (anamorph
Phyllosticta musarum (Cook) Aa). In Thailand freckle has been recorded on
various Musa species (Sontirat et al., 1994). Photita et al. (2002) recorded
G. musae Racib., G. musae Syd. & P. Syd and G. sydowiana Trotter from
Musaceae, while Photita et al. (2001) reported G. cocoicola Punith. as a common
endophyte from wild banana in northern Thailand. Brown et al. (1998) reported
P. musicola F. Stevens nom. inval. as a common endophyte from Musa acuminata
in Hong Kong. There is obviously confusion surrounding the species of these
genera occurring on banana.
The purpose of this paper is to investigate the Guignardia/Phyllosticta
spp. associated with freckle disease on leaves. We re-examined the holotype of
each epithet and also made fresh collections from banana in Asia.
MATERIAL AND METHODS
Specimens examined
Holotype specimens were loaned from S (Sweden), KRA (Poland) and
BISH (Hawaii), while fresh specimens of freckle disease on banana were collected
from Thailand. Herbarium acronyms follow Index Herbariorum (Holmgren &
Holmgren, 1998).
000_000_Wulandari.fm Page 2 Lundi, 20. décembre 2010 4:09 16

PROOF
Guignardia/Phyllosticta species on banana 3
Table 1. Countries in which banana freckle disease has been recorded
Region/country Reference Species name
Australia – New South Wales,
Queensland, Western Australia
CABI (1990),
Farr & Rossman (2010),
Jones & Alcorn (1982), Jones (1984)
Guignardia musae,
Phyllosticta musarum
Fiji Dingley et al. (1981), CABI (1990) G. musae
New Caledonia CABI (1990) G. musae
Niue Dingley et al. (1981) G. musae
Hawaii (USA) Steven (1925) G. musae
Papua New Guinea CABI (1990) G. musae
Samoa (USA) CABI (1990) G. musae
Solomon Island McKenzie & Jackson (1986),
CABI (1990)
G. musae
Tonga Dingley et al. (1981), CABI (1990) G. musae
Bangladesh CABI (1990) G. musae
Bhutan CABI (1990) G. musae
Brunei Darussalam CABI (1990),
Farr & Rossman (2010)
G. musae, P. musarum
China (Fujian, Guangdong,
Guangxi, Yunnan)
Zhou & Xie (1992),
Farr & Rossman (2010)
G. musae, P. musarum
Hong Kong, Taiwan CABI (1990) G. musae
Christmas Island Shivas & Hilton (1990) G. musae
India (Karnataka, Uttar, Pradesh) CABI (1990),
Farr & Rossman (2010)
G. musae, P. musarum
Indonesia (Java, Irian Jaya) Raciborski (1908),
Shivas et al. (1996)
G. musae
Malaysia
(Peninsular Sabah, Sarawak)
Jones (1993), CABI (1990) G. musae
Myanmar CABI (1990),
Farr & Rossman (2010)
G. musae
Nepal CABI (1990) G. musae, P. musarum
Pakistan CABI (1990) G. musae
Philippines CABI (1990) G. musae
Sri Lanka CABI (1991) G. musae
Thailand Sontirat & Jones (1994) G. musae
Vietnam Anon (1994), CABI (1990) G. musae
Cook Islands Dingley et al. (1981) G. musae
Samoa Dingley et al. (1981) G. musae
Solomon Islands McKenzie & Jackson (1986) G. musae
Vanuatu McKenzie (1989) G. musae
Palau McKenzie (1990a) G. musae
Federated States of Micronesia McKenzie (1990b) G. musae
000_000_Wulandari.fm Page 3 Lundi, 20. décembre 2010 4:09 16

PROOF
4N. F. Wulandari, C. To-Anun, L. Cai, K. A. Abd-Elsalam & K. D. Hyde
Morphology
Specimens were studied using a Nikon eclipse 80i with EOS 450 D Nikon
camera (×1000 magnification) and an Olympus CX-41 research microscope fitted
with a drawing tube and Olympus SMZ 168. Hand sections were made for
microscopic examination. Preparations and measurements were made in
lactoglycerol (lactic acid: water: glycerol = 1:2:1) for semi-permanent slide and
lactophenol cotton blue. The 95% confidence intervals were derived from
30 observations of spores formed on water agar plates, with extremes in
parentheses.
RESULTS
Taxonomy
Guignardia musae Racib., Bull. int. Acad. Sci. Lett. Cracovie, Cl. sci. math. nat.
Sér. B, sci. nat. 3: 388 (1909)
MycoBank: MB 271864
(Figs. 2-9, 17-19)
Ascomata 100-125 µm high, 75-150 µm diam, on upper and lower surface
of leaves and on banana fruit skin, globose to subglobose, black, semi-immersed
in plant tissues, coriaceous, solitary to clustered, ostiolate, ostioles as black central
dots (Fig. 2). Peridium 12.5-20 µm wide, upper part composed of compressed,
brownish, thin-walled cells, 1-4 cells thick, lower part hyaline, composed of
flattened, dark brown cells, darkest around the ostiole (Figs. 3-5, 17).
Pseudoparaphyses not observed. Asci 49-105 ×16-28 µm (›= 74 ×21 µm, n = 20),
8-spored, bitunicate, broadly cylindro-clavate, rounded at the apex, where the
diameter is 8-21 µm, tapering gradually to a 5-10 µm diam. ×5-10 µm long pedicel
attached to the basal peridium, ocular chamber 3-8 µm high (Figs. 6-7, 18).
Ascospores 20-25 ×8-13 µm (›= 22 ×10 µm, n = 20), uniseriate or occasionally
overlapping biseriate, clavate to oblong, not laterally compressed, having the same
shape when viewed from above or from the side, hyaline to greenish, 1-celled,
guttulate, smooth-walled, lacking a mucilaginous sheath or appendages at the
ends (possibly due to nature of old specimens) (Figs. 8-9, 19).
Material examined. INDONESIA, Bogor, on leaves of Musa acuminata, no date,
Raciborski, (KRA 063561, holotype of Guignardia musae Racib.), only teleomorph
present.
Notes: This is the earliest species of Guignardia or Phyllosticta described
from Musa species and therefore takes precedence over G. musae F. Stevens and
G. musae Syd. & P. Syd. The ascospores in this species are distinct because of
their size (20-25 ×8-13 µm) and shape (clavate to oblong, not laterally compressed
having the same shape when viewed from above or the side) (Table 2).
Guignardia stevensii Wulandari & K.D. Hyde, nom. nov.
!Guignardia musae F. Stevens, Bulletin of the Bernice P. Bishop Museum,
Honolulu, Hawaii 19: 101 (1925), nom. illegit., non G. musae Racib. 1909.
MycoBank: MB 519089
(Figs. 10-13, 20-22)
Etymology: Named after its collector, F.L. Stevens.
000_000_Wulandari.fm Page 4 Lundi, 20. décembre 2010 4:09 16

PROOF
Guignardia/Phyllosticta species on banana 5
Figs. 1a-d. Freckle disease on Musa spp. in Thailand caused by Guignardia musicola.
000_000_Wulandari.fm Page 5 Lundi, 20. décembre 2010 4:09 16

PROOF
6N. F. Wulandari, C. To-Anun, L. Cai, K. A. Abd-Elsalam & K. D. Hyde
Figs. 2-16. Micrographs of Guignardia spp. on Musa sp. 2-9. G. musae Racib., 10-13. G. stevensii,
15-16. G. musicola.2. Appearance of ascomata on host surface (bar = 100 µm). 3, 4, 5. Section of
ascoma in the leaf (darkened area fungal cells-arrowed) (bar = 20 µm). 6, 7. Asci (bars 6 = 30 µm,
7 = 10 µm). 8, 9. Ascospores (bar = 10 µm). 10, 11. Asci (bar = 15 µm). 12, 13. Ascospores
(bar = 5 µm). 14. Asci (bar = 10 µm). 15, 16. Ascospores (bar = 10 µm).
000_000_Wulandari.fm Page 6 Lundi, 20. décembre 2010 4:09 16

PROOF
Guignardia/Phyllosticta species on banana 7
Figs. 17-19. Line drawing of G. musae Racib. (holotype): 17. Section of ascoma in the leaf (fungal
cells arrowed) (bar = 25 µm). 18. Asci (bar = 25 µm). 19. Ascospores which are clavate to oblong
and symmetrical (bar = 25 µm).
000_000_Wulandari.fm Page 7 Lundi, 20. décembre 2010 4:09 16

PROOF
8N. F. Wulandari, C. To-Anun, L. Cai, K. A. Abd-Elsalam & K. D. Hyde
Ascomata 50-125 µm high, 60-95 µm diam, on upper surface of leaves,
globose to subglobose, black, semi-immersed in plant tissues, coriaceous, solitary
to clustered, ostiolate, ostioles as black central dots. Peridium 20-25 µm wide,
composed of compressed, brownish, thin-walled cells, in the upper part 1-4 cells
thick, composed of flattened, dark brown cells, darkest around the ostiole, hyaline
towards the lower region (Fig. 20). Pseudoparaphyses not observed. Asci
40-59 ×11-15 µm (›= 50 ×13 µm, n = 20), 8-spored, bitunicate, cylindro-clavate,
rounded at the apex, where the diameter is 10-12 µm, tapering gradually to a
2-7 µm diam. ×3-7 µm long pedicel attached to the basal peridium, ocular
chamber 3-8 µm high (Figs. 10-11, 21). Ascospores 14-17 ×5-6 µm (›= 15 ×5µm,
n=20),uniseriateoroccasionallyoverlappingbiseriate,ellipsoidal,widest2/5
th from
the apex (obtrullate) in one plane, inequilaterally ellipsoidal, or ellipsoidal with one
side flattened when viewed from the side, hyaline to greenish, 1-celled, guttulate,
smooth-walled, with a mucilaginous appendage at each end (Figs. 12-13, 22).
Material examined. HAWAII, Oahu, Hakipu, on leaves of Musa sp., 12 June 1921,
F.L. Stevens, No. 565 (BISH 596860, holotype; BISH 499904 isotype of Guignardia musae
F. Stevens), teleomorph only present.
Notes: The ascospores of Guignardia musae F. Stevens differ markedly
from those of G. musae Racib. being 14-17 ×5-6 µm, obtrullate from above,
inequilaterally ellipsoidal, or ellipsoidal and flattened on one when viewed from
Table 2. Synopsis of ascospores and conidia of Guignardia species on Musa spp.
G. musae Racib. G. stevensii G. musicola
Ascus (µm) 49-105 ×16-28,
broadly clavate
40-59 ×11-15,
cylindro-clavate
133-150 ×19-20, cylindrical
to cylindro-clavate
Ascospores (µm) 20-25 ×8-13, clavate
to oblong symmetrical,
without appendages
14-17 ×5-6, widest 2/5 near
the apex (obtrullate),
inequilateral, or ellipsoidal
with one side flattened,
and with appendages
12-21 ×7-10, obclavate
to oblong, symmetrical,
with appendages
Line drawing
of ascospores, bars
(a = 25 µm;
b-c = 20 µm)
Phyllosticta state
(µm)
Not present Not present Conidia 12-17 ×8-11,
with appendage 10-15 long,
sheath 2-4 wide
000_000_Wulandari.fm Page 8 Lundi, 20. décembre 2010 4:09 16

PROOF
Guignardia/Phyllosticta species on banana 9
Figs. 20-22. Line drawing of G. stevensii (holotype): 20. Section of ascoma in the leaf (fungal cells
arrowed) (bar = 25 µm). 21. Asci (bar = 20 µm). 22. Ascospores which are obtrullate from above,
inequilaterally ellipsoidal, or ellipsoidal with one side flattened, and with mucilaginous
appendages at the ends (bar = 20 µm).
000_000_Wulandari.fm Page 9 Lundi, 20. décembre 2010 4:09 16

PROOF
10 N. F. Wulandari, C. To-Anun, L. Cai, K. A. Abd-Elsalam & K. D. Hyde
the side (Table 2). Since G. Musae F. Stevens is a homonym of G. Musae Racib.
we provide a new name. Fresh living collections from Hawaii are needed to fully
circumscribe this taxon from Musa sp. with DNA sequence comparison.
Guignardia sydowiana Trotter, in Saccardo, Syll. Fung. (Abellini) 24(2): 788
(1928)
Basionym. Guignardia musae Syd. & P. Syd., Annls mycol. 10: 80 (1912)
[name is invalid as homonym of G. musae Racib.].
Material examined. Democratic Republic of Congo, on dead leaf of Musa sp., Vanderyst, ex
Herb. Sydow (S, 10753, holotype of Guignardia musae Syd. & P. Syd).
Notes: The name Guignardia sydowiana Trotter was introduced to
replace G. musae Syd. & P. Syd., which is a homonym of G. musae Racib., and
thus invalid. The type material examined is not in a good condition as ascomata
were dry and depauperate.
Guignardia musicola N.F. Wulandari, L. Cai & K.D. Hyde, sp. nov.
MycoBank no.: MB 519088
(Figs. 14-16, 23-31)
Etymology: Named after its host plant, Musa sp. and -cola meaning dwelling on.
Guignardiae musae Racib. similis,sed ascosporae 12-21 ×7-10 µm.
Leaf spot occupying marginal areas of the leaf and pinna, bleached, the
leaf breaking at the edge to the middle of lamina, ascomata visible to the unaided
eye on surface of the leaves, surface rough indicating protruding ascomata and
pycnidia (Fig. 1). Ascomata 100-125 µm diam, 100-125 µm high, on upper surface
of leaves, globose to subglobose, black, semi-immersed in plant tissues,
coriaceous, solitary to clustered, ostiolate, ostioles as black central dots. Peridium
22.5-25 µm wide, comprising 2 layers of textura angularis cells with thickened
brown walls around ostiole (Fig. 23). Pseudoparaphyses not observed. Asci
133-150 ×19-20 µm (›= 137 ×20 µm, n = 20), 8-spored, bitunicate, fissitunicate,
cylindrical to cylindro-clavate, rounded at the apex, where the diameter is
12-13 µm, tapering gradually to a 10-20 µm diam. ×5-6 µm long pedicel attached
to the basal peridium, ocular chamber 2-5 µm high (Figs. 14, 24). Ascospores
12-21 ×7-10 µm (›= 19 ×9µm, n = 20), uniseriate or occasionally overlapping
biseriate, ellipsoidal to clavate, not laterally compressed, having the same shape
when viewed from above or the side, hyaline to greenish, 1-celled, guttulate,
smooth-walled, with a mucilaginous appendage at each end, not (Figs. 15-16, 25).
Pycnidia 95-125 µm diam, 75-125 µm high, epiphyllous, black, globose to pyriform,
immersed in plant tissues, coriaceous, solitary to clustered, ostiolate, ostioles as
white dots in the centre. Peridium 22-25 µm wide, one stratum of textura angularis
comprising 2 layers of cells with thickened brown walls around ostiole (Fig. 26).
Conidiogenous cells 10-12 ×8-9 µm (›= 11 ×8 µm, n = 5), holoblastic,
determinate, discrete, sometimes rarely integrated, hyaline, cylindrical to
doliiform cells lining the pycnidial locule (Fig. 27). Conidia 12-17 ×8-11 µm
(›= 14 ×10 µm, n = 20), hyaline to greenish, 1-celled, guttulate, smooth-walled,
globose, ellipsoidal, clavate or obclavate, with an obtuse apex, sometimes truncate
at the base, surrounded by 2-4 µm thick mucilaginous sheath which persists at
maturity and in some specimens with a single, hyaline, curved or straight,
10-15 µm long appendage (Fig. 28). Spermogonia 95-125 µm in diameter,
75-125 µm high, epiphyllous, black, globose to subglobose, immersed in plant
tissues, coriaceous, solitary to clustered, ostiolate, ostioles as white dots in the
centre, similar to pycnidia. Peridium 22-25 µm wide, one stratum of textura
000_000_Wulandari.fm Page 10 Lundi, 20. décembre 2010 4:09 16

PROOF
Guignardia/Phyllosticta species on banana 11
Figs. 23-25. Line drawing of G. musicola (holotype): 23. Section of ascoma in the leaf (fungal cells
arrowed) (bar = 25 µm). 24. Asci (bar = 20 µm). 25. Ascospores which are obclavate to oblong,
symmetrical, with appendages (bar = 20 µm).
000_000_Wulandari.fm Page 11 Lundi, 20. décembre 2010 4:09 16

PROOF
12 N. F. Wulandari, C. To-Anun, L. Cai, K. A. Abd-Elsalam & K. D. Hyde
Figs. 26-28. Line drawing of Phyllosticta state of G. musicola (holotype): 26. Section of pycnidium
in the leaf (fungal cells arrowed) (bar = 10 µm). 27. Conidia and conidiogenous cells
(bar = 10 µm). 28. Conidia (bar = 10 µm).
000_000_Wulandari.fm Page 12 Lundi, 20. décembre 2010 4:09 16

PROOF
Guignardia/Phyllosticta species on banana 13
Figs. 29-31. Line drawing of Leptodothiorella state of G. musicola (holotype): 29. Section
of spermogonium in the leaf (fungal cells arrowed) (bar = 25 µm). 30. Spermatiogenous cells
(bar = 10 µm). 31. Spermatia (bar = 10 µm).
000_000_Wulandari.fm Page 13 Lundi, 20. décembre 2010 4:09 16

PROOF
14 N. F. Wulandari, C. To-Anun, L. Cai, K. A. Abd-Elsalam & K. D. Hyde
angularis comprising 2 layers of cells with thickened brown walls around ostiole
(Fig. 29). Spermatiogenous cells 7-10 ×1 µm (›= 9.8 ×1 µm, n = 20), holoblastic,
filamentous to cylindrical, simple or branched as distinct phialides with a very
characteristic and easily discernible apical structure (Fig. 30). Spermatia 5-8 ×1-2 µm
(›=7×1µm, n = 20), cylindrical to dumb-bell shaped, guttulate, straight or
slightly curved forming singly in basipetal succession and separating from the
spermatiogenous cells by a septum (Fig. 31).
Material examined. THAILAND, Chiang Mai Province, Chiang Mai, Tung Jaow Village, on
leaves of Musa acuminata, 18 July 2007, N.F. Wulandari, NFW 154 (MFLU 10 0235,
holotype) teleomorph and anamorph present; extype cultures CBS 123405; ibid., Srilanna,
on leaves of Musa paradisiaca, 12 July 2007, N.F. Wulandari, NFW 140 (MFLU 10 0233)
teleomorph and anamorph present; Bahn Pa Deng, T. Pa Pae, Mae Taeng, Mushroom
Research Centre, on leaves of M. paradisiaca, 24 August 2006, N.F. Wulandari, NFW 084
(MFLU 10 0222), teleomorph only present; ibid., 3 June 2007, N.F. Wulandari NFW 128
(MFLU 10 0231), teleomorph only present; ibid., 20 July 2007, N.F. Wulandari, NFW 161
(MFLU 10 0236), teleomorph only present; ibid., 13 August 2007, N.F. Wulandari, NFW 176
(MFLU 10 0237), teleomorph and anamorph present; ibid., 21 August 2007, N.F. Wulandari,
NFW 182 (MFLU 10 0238), teleomorph and anamorph present; ibid., 12 September 2007,
N.F. Wulandari, NFW 219 (MFLU 10 0244), teleomorph only present. Tumbon,
Chiangdoaw, on leaves of M. paradisiaca, 5 September 2007, N.F. Wulandari, NFW 184
(MFLU 10 0239), teleomorph, anamorph and spermatial stage present; ibid., 5 September
2007, N.F. Wulandari, NFW 185 (MFLU 10 0240), teleomorph only present; ibid.,
5 September 2007, N.F. Wulandari, NFW 188 (MFLU 10 0242), teleomorph only present.
Bahn Pha Deng, Mae Lod, Royal Project, on leaves of M. paradisiaca, 11 September 2007,
N.F. Wulandari, NFW 210 (MFLU 10 0243), teleomorph only present; ibid., Chiang Mai,
Chiang Mai University on leaves of M. paradisiaca, 16 June 2006, N.F. Wulandari, NFW 114
(MFLU 10 0225), teleomorph only present; ibid., 19 June 2007, N.F. Wulandari, NFW 117
(MFLU 10 0228), teleomorph only present; ibid., 19 June 2006, N.F. Wulandari, NFW 118
(MFLU 10 0229), teleomorph only present; Chiang Mai University Shop garden, on leaves
of M. paradisiaca, 15 September 2007, W. Tajeena & N.F. Wulandari, NFW 221
(MFLU 10 0245), teleomorph and anamorph present; Medicinal Plant Garden on leaves of
Musa paradisiaca, 15 September 2007, N.F. Wulandari, NFW 230 (MFLU 10 0246),
teleomorph only present. Bahn Pha Deng, Pathummikaram Temple, on leaves of
M. paradisiaca, 1 July 2007, N.F. Wulandari, NFW 123 (MFLU 10 0230), teleomorph only
present; Bahn Pha Deng Mushroom Research Centre, on leaves of M. paradisiaca,
24 August 2006, N.F. Wulandari, NFW 079 (MFLU 10 0220), teleomorph only present; ibid.,
22 August 2006, N.F. Wulandari NFW 080 (MFLU 10 0221), teleomorph only present; ibid.,
18 June 2007, N.F. Wulandari, NFW 115 (MFLU 10 0226), teleomorph only present;
ibid., 18 June 2007, N.F. Wulandari, NFW 116 (MFLU 10 0227), teleomorph only present; ibid.,
N.F. Wulandari, NFW 131 (MFLU 10 0232), teleomorph only present; ibid., 17 July 2007,
N.F. Wulandari NFW 151 (MFLU 10 0234), teleomorph only present. Chiang Rai, Nam Tok
Huey Mesak Forest Park, on leaves of M. paradisiaca, 6 February 2010, N.F. Wulandari &
P. Syshophanthong, NFW 306 (MFLU 10 0281), teleomorph only present.
Notes: Guignardia musicola is distinct from G. musae Racib. as
ascospores in G. musicola are smaller 12-21 ×7-10 µm (›=19×9µm), compared
with those of Guignardia musae Racib. (20-25 ×8-13 µm, ›=22×10 µm) (Table 2).
DISCUSSION
This study redescribes G. musae Racib. and shows it to be a
morphologically distinct species. Fresh collections are needed from Indonesia,
however, to epitypify this species for molecular study. Guignardia musae
000_000_Wulandari.fm Page 14 Lundi, 20. décembre 2010 4:09 16

PROOF
Guignardia/Phyllosticta species on banana 15
F. Stevens and G. musae Syd. & Syd. are homonyms of G. musae Racib. and thus
invalid. Guignardia musae F. Stevens is, however, a distinct species and a new
name G. stephensii is introduced for this taxon . One new species of Guignardia
isolated from leaves of banana with freckle symptoms in Thailand (Fig. 1) is also
introduced. The study shows that more than one species is responsible for freckle
symptoms of banana and a worldwide study is justified. Several other species, e.g.
Macrophoma musae (Sacc.) Berl. & Voglino, Phoma musae Sacc., Phoma musae
C.W. Carp., Phyllosticta musarum (Cooke) Aa, Sphaeropsis musarum Cooke and
Phyllostictina musarum (Cooke) Petr. have at one time or another been
considered to be synonyms of G. musae Racib. (Aa, 1973; Carpenter, 1919; Petrak
and Ciferri, 1931; Raciborski, 1909; Sivanesan, 1984). The synonymies, however,
were based on morphological data and the taxa need recollecting and subjecting
to molecular analysis. Futher collections and sequence analysis are needed from
different continents and various musaceous hosts to establish which species induce
freckle disease of banana.
Acknowledgements. Nilam Wulandari acknowledges the Graduate School of
Chiang Mai University, Chiang Mai, Thailand for financial support. The herbaria,
BISH, KRA and S are thanked for loaning specimens. The authors are grateful to
P. Sysouphanthong, P. Phengsintham, S. Karunarathna, and W. Tajeena who helped
collecting banana leaf samples. Mae Fah Luang University and Hong Kong University are
thanked for laboratory facilities. Shaun Pennycook is thanked for advice on the Latin
names introduced in this paper. The Mushroom Research Foundation and CBS, the
Netherlands, are thanked for a PhD scholarship. BRT, Thailand are also thanked for
awarding a grant (BRT No R251181) to study Dothideomycetes in northern Thailand.
Professor P. Crous is thanked for partially funding this research. The authors also gratefully
acknowledge partial financial support from the Distinguished Scientist Fellowship Program
(DSFP) King Saud University. Thida Win Ko KO and Eric McKenzie are gratefully
acknowledged for providing suggestions to improve the draft manuscript.
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000_000_Wulandari.fm Page 16 Lundi, 20. décembre 2010 4:09 16

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6(2)D*]*K"C*5)$3*Morinda citrifolia9* X'=7(*<1*
H/RS+I* 2)14%5())(C* =2* 1%* Phyllosticta morindae*
H6(2)D* ]* K"CDI* <1* 14C* 7=4N(C* =2* X=2'* 2'(*
2(7($3$)&'* GD morindaeD* :1))* ]* F$%%314*
H,A/AI*7=%2* GD morindae* 5)$3* <;%2)17=19*J4C=19*
J4C$4(%=19* M1&14* 14C* K13$19* E;2* =2* =%* @()"*
#$33$4* =4* 61#=5=#* =%714C* #$;42)=(%* $5*
<3()=#14* K13$19* >$$N* J%714C%9* :(C()12(C*
K212(%* $5* !=#)$4(%=19* :=c=9* :)(4#'* 6$7"4(%=19*
O=)=E12=9* U=;(9* 6171;9* K13$19* B$4819* B;@17;9*
a14;12;9* T177=%* 14C* :;2;41* H0=487("* (2* 17D*
/R_/9* !#O(4b=(* /R_R9* /RRA19E9* /RRQ9* K'=@1%*
/RRQID*
Morinda citrifolia*HF;E=1#(1(I* =%*1*'=8'V
7"*&)=b(C*3(C=#=417*&7142D* U$4=*c;=#(*(Y2)1#2(C*
5)$3*2'(*5()3(42(C*5);=2*=%*31)N(2(C*X$)7CX=C(*
14C* '1%* 14* (%2=312(C* @17;(* $5* ?Kd,* E=77=$4*
144;177"D*!$%2*$5*2'(*)1X*&)$C;#2*#$3(%*5)$3*
6$7"4(%=1D* >);C(* (Y2)1#2%* $5* MD citrifolia*14C*
MD elliptica* LD* '1@(* E((4* %'$X4* 2$* '1@(*
142=@=)17* 1#2=@=2"* 181=4%2* 5$$2* 14C* 3$;2'*
C=%(1%(* @=);%* H>';48%131)4"1)2* (2* 17D* ,AASID*

+,Q*
e(4$2$Y=#* 14C* 142=8(4$2$Y=#* (55(#2%* $5* 4$4=*
5);=2*c;=#(*&)$C;#(C*=4*B'1=714C* '1C*8(4$2$Y=#*
14C* 142=8(4$2$Y=* (55(#2%* $4* ';314* 7"3&'$V
#"2(%*=4* 2'(* #')$3$%$3(*1E())12=$4* 1%%1"*14C*
%=%2()*#')$312=C*(Y#'148(*HK>fI*1%%1"*in vitroD*
HF12141@171#'1=*(2*17D*,AA_9*B'14=*(2*17D*,A/AID*
U$4=* 1&&(1)%* 2$* )(%2$)(* 2'(* 4$)317* 3(4%2);17*
#"#7(* &)$E7(3%* 14C* 177(@=12(* 3(4%2);17* %"3&V
2$3%*=4*3=#(*H>'(1)%N;7*(2*17D*,AAP9*B'14=*(2*17D*
,A/AI*14C*=4'=E=2%*3;)=4(*2;3$)*8)$X2'*X=2'*1*
C(5=4=2(* #;)12=@(* &$2(42=17* =4* 3=#(* H:;);%1X1*
,AA,ID*!12'=@1414* (2*17D*H,AA-I* )(@=(X(C*#;)V
)(42* )(%(1)#'* $4* Morinda citrifolia*X'=7(*
F(2'=413* ]* K=@1)1314* H,AASI* C=%#;%%(C*
)(%(1)#'* C(@(7$&3(42%* =4* J4C=1* 14C* (7%(X'()(*
14C* )(@=(X(C* 2'(* 7=2()12;)(D* B'(* $Ec(#2=@(* $5*
2'(*)(%(1)#'*=%*2X$*5$7CD*B'=%*&1&()*&)$@=C(%*14*
;&C12(C*C(%#)=&2=$4*$5*GD Morindae*14C9*%=4#(*
2'(*2"&(*%&(#=3(4*=%*7$%2*1*4($2"&(*=%* C(%=841V
2(C*=4* $)C()*2$* %21E=7=b(* 2'(*1&&7=#12=$4* $5*2'=%*
%&(#=(%*413(D*
*
Results
>$77(#2=$4%* $5* Guignardia morindae
5)$3*2')((*C=55()(42*7$#12=$4%*1)(*#$3&1)(CD* <*
C(%#)=&2=$4* 14C* =77;%2)12=$4* 5)$3* 2'(* 4($2"&(*
%&(#=3(4*=%* 31C(*14C* 1*4($2"&(* =%*C(%=8412(C*
'()(D*
*
Taxonomy
*
Guignardia morindae HO$$)CDI* <19* StudD
MycolD*-[*QR*H/RS+I* :=8%*/.++
!"#$G14N*+/PS-Q*
Ł*Physalospora morindae* O$$)CD9* VerhD
KD AkadD VetD Amsterdam*/+HPI[*/RA*H/RASID*
Ł*Puiggarina morindae*HO$$)CDI*K&(8D9*
Boletín de la Academia Nacional de Ciencias
de Córdoba*,+[*P_Q*H/R/RID*
<413$)&'* Phyllosticta morindae*H6(2)D*
]*K"CDI*<19*StudD MycolD -[*QR*H/RS+ID**
Ł*Phyllostictina morindae* 6(2)D* ]* K"CD9*
Feddes RepertD9*G(='D*P,[*,AA*H/R,SID*
>1;%=48*5)$8*("(* $)*%'$2V'$7(*7(15* %&$2%9*
AD+.AD_* g* AD-./D,* #39* X'=#'* 1)(* )$;4C(C* 2$*
=))(8;71)* X=2'* )(C* 2$* C1)N* E)$X4* E$)C()%h* 2'(*
1)(1* X'()(* 2'(* 5;48=* %&$);712(%* =%* 2)14%&1)(42*
14C* $52(4* 5177%* 5)$3* 2'(* 7(15* H:=8%* /9* ,9* +ID*
<%#$3121* QA./,A* i3* C=13(2()9* RA./AA* i3*
'=8'9* $4* 2'(* ;&&()* 14C* 7$X()* %;)51#(%* $5* 2'(*
7(1@(%9*E71#N9*87$E$%(*2$*%;E87$E$%(9*=33()%(C*
=4*&7142*2=%%;(%9*#$)=1#($;%9*#7;%2()(C9*$%2=$712(9
*
Figs 1.5 .* Guignardia*5)$8*("(*C=%(1%(*$4*
7(1@(%D* 1.3* K"3&2$3%* $4* 7(1@(%D* 4.5*
<&&(1)14#(*$5*1%#$31*$4*2'(*'$%2*%;)51#(D*
*
$%2=$7(%*1%*E71#N*C$2%*=4*2'(*#(42)(*H:=8%*P9*-9*Q9*
S9* _ID* 6()=C=;3* /A./-* i3* X=C(9* #$3&$%(C* $5*
2X$*2$*2')((*71"()%*$5*#(77%9*$5*textura angularis9*
&=83(42(C* $;2X1)C7"* 14C* 1)$;4C* $%2=$7(* 14C*
&17()*=4%=C(*H:=8%*R9*/A9*/QID*6%(;C$&1)1&'"%(%*
'"&'1V7=N(9*,.+*ȝ3*=4*C=13D*<%#=*+R.Q-*g*//.
/P* i3* H
x
*j* -A* g* /,* i39* 4* j* ,AI9* _V%&$)(C9*
E=2;4=#12(9* 5=%%=2;4=#12(9* E)$1C7"* #"7=4C)$V
#71@12(9*)$;4C(C*12*2'(*1&(Y9*21&()=48*8)1C;177"*
2$* 1* &(C=#(7* 1221#'(C* 2$* 2'(* E1%17* &()=C=;3*
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5)$3* 1E$@(* 14C* =4(k;=712()177"* (77=&%$=C17* $)*
(77=&%$=C17* X=2'* $4(* %=C(* 57122(4(C* C$)%177"*
X'(4*@=(X(C*5)$3*%=C(9*'"17=4(*$)*8)((4=%'9*/V
#(77(C9* #$1)%(V8;22;712(9* %3$$2'VX177(C9* X=2'*
)$;4C(C* (7$4812(* (4C%* 14C* E=&$71)* 3;#=71V
8=4$;%*1&&(4C18(%*H:=8%*/,./-9*/RID**
6"#4=C=1*_-.R-* i3*C=13(2()9* QP._-*i3*
'=8'9*$4*2'(*;&&()*14C*7$X()*7(15*%;)51#(9*E71#N9*
87$E$%(* 2$* %;E87$E$%(9* =33()%(C* =4* &7142* 2=%V
%;(%9*#$)=1#($;%9*%$7=21)"*2$*#7;%2()(C9*$%2=$712(9

!"#$%&'()(*
**
+,S*
*
Figs 6.15 .*Guignardia morindae H4($2"&(I. 6 L(15*%&$2%D*7, 8*<&&(1)14#(*$5*1%#$3121*$4*2'(*'$%2*
%;)51#(D*9 K(#2=$4*$5*1%#$31D*10*6()=C=;3*#$3&)=%=48*$4(*%2)121*$5*2(Y2;)1*148;71)=%*#$3&)=%=48*,.
+*71"()%*$5*#(77%*X=2'* 14*1&(Y*$5*2'=#N(4(C*E)$X4* X177%D*11 <%#=D*12, 13, 14, 15*<%#$%&$)(%*X=2'*
E=&$71)*3;#=718=4$;%*1&&(4C18(%*X=2'*)$;4C(C*(7$4812(*(4C%*.*G1)%*/,*j*-A*i39*/+*j*,A*i39*/P*j*
/A*i39*/-./_*j*-*i3D*
*
$%2=$7(%* 1%* E71#N* C$2%* =4* 2'(* #(42)(9* $52(4*
8)$X=48*2$8(2'()*X=2'*1%#$3121D*6()=C=;3*//.
/-*i3* X=C(9* #$3&$%(C* $5*2X$* 2$* 2')((*71"()%*
$5* #(77%9* textura angularis* 14C* &=83(42(C*
$;2X1)C7"* 14C* 1)$;4C* $%2=$7(* 14C* &17()* =4%=C(*
H:=8%*,A9* ,/9*,_ID* >$4=C=$8(4$;%* #(77%*S./,* g*
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x
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7=4=48* 2'(* &"#4=C=17* 7$#;7(* H:=8%* ,A9* ,RID*
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$E#71@12(9* X=2'* 14* $E2;%(* 1&(Y9* %$3(2=3(%*
2);4#12(*$4* 2'(* E1%(9*%;))$;4C(C* E"*AD-./* ȝ3*
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x
j*/* i39* 4* j*,AI* 2'=#N* 3;#=718=4$;%*%'(12'*
X'=#'*&()%=%2%*12*312;)=2"*X=2'*1*,.S*ȝ3*H
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K&()3$8$4=1*PP.P-*i3*C=13(2()9*P,.PS*
i3*'=8'*=42()3=Y(C*X=2'*&"#4=C=1D*6()=C=;3*-.
R*i3*X=C(9*#$3&$%(C*$5*2X$*2$*2')((*71"()%*$5*
#(77%9* textura angularis*14C*&=83(42(C*$;2V
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x
j*/Q*g*,*i39*4*j*,AI9*'$7$E71%2=#9*
5=713(42$;%* 2$* #"7=4C)=#179* %=3&7(* $)* E)14#'(C*
14C*(1%=7"*C=%#()4=E7(*1&=#17*%2);#2;)(*H:=8%*,Q9*
+,ID*K&()312=1*-.R*g*/.,*i3*H
x
j*Q*g*,*i39*4*
j* ,AI* '$7$E71%2=#9* #"7=4C)=#17* 2$* C;3EVE(7*
%'1&(C9* 8;22;712(9* %2)1=8'2* $)* %7=8'27"* #;)@(C*
5$)3=48* %=487"* =4* E1%=&(217* %;##(%%=$4* 14C

+,_*
*
Figs 16.19 .* Guignardia morindae H4($2"&(I*
7=4(*C)1X=48D*16 K(#2=$4*$5*1%#$31D*17 !12;)(
<%#=D*18*J3312;)(*<%#=D*19*<%#$%&$)(%D**
*
%(&1)12=48*5)$3*2'(*%&()312=$8(4$;%*#(77%*E"*1*
%(&2;3*H:=8%*,S9*++ID*
!12()=17*(Y13=4(C*.*JU0`UfKJ<9*T(%2*
M1@1*6)$@=4#(9*G$8$)9*G$8$)*G$214=#17*e1)C(49*
$4* 7=@=48* 7(15* $5* Morinda citrifolia9 ,+*
K(&2(3E()* ,A/A9* UD:D* T;714C1)=9* U:T* +Q/*
HG`*,,QP_* C(%=8412(C* 1%*4($2"&(I9* %&()312=17*
%218(9*1413$)&'*14C*2(7($3$)&'*&)(%(42h*ibidD9*
,+*K(&2(3E()*,A/A9*UD:D*T;714C1)=9*U:T*+Q+*
HG`*,,Q-AI9*2(7($3$)&'*$47"*&)(%(42h*ibidD9*//*
!1"* ,AAQ9* UD:D* T;714C1)=9* U:T* /QR* HG`*
,,Q-,I9* 2(7($3$)&'* 14C* 1413$)&'* &)(%(42h*
ibidD9*,S*M;4(*,AAQ9*UD:D*T;714C1)=9*U:T*/Q_*
HG`* ,,Q-/I9* 1413$)&'* $47"* &)(%(42h* >(42)17*
M1@1* 6)$@=4#(9* U8(421N9* U8(421N9* O(C;9* $4*
7=@=48*7(15* $5*Morinda citrifolia*/_* K(&2(3E()*
,A/A9*UD:D* T;714C1)=9*U:T*+Q,* HG`*,,QPRI9*
1413$)&'* $47"* &)(%(42D* BW<JL<U09* >'=148*
F1=9*6'1'$4"$2'=4*F$1C9*$4*7(1@(%*$5*Morinda
citrifolia9* ,A* M14;1)"* ,A/A9* UD:D* T;714C1)=9*
U:T* ,RQ* H!:L?* /A* AP-+I9* 2(7($3$)&'* $47"*
&)(%(42h*ibidD9*A-*!1)#'*,A/A9*UD:D*T;714C1)=9*
U:T* +/+* H!:L?* /A* APQQI9* 2(7($3$)&'* $47"*
&)(%(42D*
O4$X4* C=%2)=E;2=$4* .* <3()=#14* K13$19*
<;%2)17=19* >$$N* J%714C%9* :(C()12(C* K212(%* $5*
!=#)$4(%=19*:=c=9*:)(4#'* 6$7"4(%=19*T177=%*14C*
:;2;419*J4C=19*J4C$4(%=1*HG$8$)9*O(C;I9*M1&149
*
*
Figs 20.27 .*Phyllosticta %212(*$5*G. morindae*
H4($2"&(ID*20, 219*24, 25*6()=C=;3*#$3&)=%=48*
$4(*%2)121*$5*textura angularis*#$3&)=%=48*,.+*
71"()%*$5*#(77%*X=2'*14*1&(Y*$5*2'=#N(4(C*E)$X4*
X177%D*22 >$4=C=$8(4$;%* #(77%D*23*>$4=C=1D*24
K&()3$8$4=1D 26 K&()312=$8(4$;%* #(77%* 27*
K&()312=1*.*G1)%*,A*j*P-*i39*,/*j*/-*i39*,,*
j*+*i39*,+*j*S*i39*,P*j*,A*i39*,-9*,S*j*R*i39*
,Q*j*,,*i3D*
*
O=)=E12=9U=;(9*6171;9*K131$9*B'1=714C* H>'=148*
!1=9*>'=148*F1=I9*B$4819*B;@17;9*a14;12;*14C*
T177=%D**
*
Discussion
B'(*'$7$2"&(*$5* Guignardia morindae*=%*
4$2* =4* G`h* O$$)C()%* 4(@()* C(&$%=2(C* '=%*
%&(#=3(4%* =4* 2'(* '()E1)=;3* H!=(4* <D* F=51=9*
&()%D*#$33DI*14C* 2'()(*=%*4$* 1@1=71E7(*(YV2"&(*
#;72;)(D*K=4#(* 2'()(*=%*4$* 2"&(*$5* GD morindae*
2'(* %&(#=(%* X1%* )(#$77(#2(C* 5)$3* G$8$)* 12* 2'(

!"#$%&'()(*
**
+,R*
*
*
*
Figs 28.30 .*Phyllosticta %212(*$5*G. morindae
H4($2"&(I*7=4(*C)1X=48D*28 K(#2=$4*$5*&"4=C=;3D*
29 >$4=C=$8(4$;%*#(77%D*30*>$4=C=1D**
*
$)=8=417* &71#(* $5* #$77(#2=$4D* <* 4($2"&(* =%* 1*
%&(#=3(4* $)* =77;%2)12=$4* %(7(#2(C* 2$* %()@(* 1%*
4$3(4#712;)17* 2"&(* =5* 4$* $)=8=417* 312()=17* =%*
(Y21429* $)* =%* 3=%%=48* H<)2* RDQI* H!#U(=77* (2* 17D*
,AAQID*B'(*4((C*$5*4($2"&=5=#12=$4*=%*=3&$)2142*
=4* $)C()* 2$* %21E=7=b(* 2'(* 1&&7=#12=$4* $5* 2'(*
%&(#=(%* 413(* H!#U(=77* (2* 17D* ,AAQID* Guignar-
dia morindae* =%* )(#$)C(C* 5$)* 2'(* 5=)%2* 2=3(* =4*
B'1=714CD*!$7(#;71)*X$)N*=%*4((C(C* 2$*C=%#()4*
2'(* C=%2=4#24(%%* $5* 2'=%* %&(#=(%D* W$X(@()9*
C(%&=2(* )(&(12(C* 122(3&%* X(* #$;7C* 4$2* =%$712(*
2'(*5;48;%D*
*
Acknowledgements
U=713*T;714C1)=*=%*8)12(5;7*2$*2'(*e)1CV
;12(* K#'$$79* >'=148* !1=* ?4=@()%=2"9* B'1=714C*
5$)*5=414#=17*%;&&$)2*14C*2'(*K#'$$7*$5*K#=(4#(9*
!1(* :1'* L;148* ?4=@()%=2"* 5$)* 71E$)12$)"*
51#=7=2=(%D* M$%%(* F=b17* =%* 2'14N(C* 5$)* 5=414#=17*
%;&&$)2* 5$)* 2'(* #$77(#2=48* 2)=&* =4* G$8$)9* T(%2*
M1@1*14C* O(C;9* >(42)17*M1@19* J4C$4(%=1D*?E(4*
K"1)=5;CC=49* F=%3=21* K1)=* 14C* J)1* X12=9* G$8$)*
G$214=#17* e1)C(49* G$8$)9* J4C$4(%=1* 14C* K)=*
U812=4=9* U8(421N9* O(C;9* >(42)17* M1@19*
J4C$4(%=1* 1)(* 2'14N(C* 5$)* Morinda citrifolia*
7(15* %13&7(%D* W()E1)=;3* G$8$)=(4%(* HG`I9*
J4C$4(%=1*2')$;8'*fN$*G1)$2$*T17;c$9*O1)2=4=**
Figs 31.33 .* Leptodothiorella %212(* $5* G.
morindae H4($2"&(I*7=4(*C)1X=48D*31 K(#2=$4*$5*
%&()3$8$4=;3D* 32 K&()312=$8(4$;%* #(77%D* 33*
K&()312=1D*
*
O)131C=E)121* 1)(* 2'14N(C* 5$)* %&(#=3(4%* #()2=*
5=#12(* &()3=2* U$D* //_PlJ6WD/DA,lODKDA/DAPl*
,A/AD* !=(4* <D* F=51=* 14C* 0(X=* K;%14* 1)(*
2'14N(C* 5$)* @17;1E7(* =45$)312=$4* #$4#()4=48*
2'(*'$7$2"&(*$5*Physalospora morindae =4*G`D*
!1(* :1'* L;148* ?4=@()%=2"* =%* 2'14N(C* 5$)* 2'(*
1X1)C*$5*1*8)142*4$*-+/A/A,AA/S*H/Sl,--+I*2$*
%2;C"* Phyllosticta =4* 4$)2'()4* B'1=714CD* GFB9*
B'1=714C* =%* 1#N4$X7(C8(C*5$)*2'(*1X1)C*$5*1*
8)142* HGFB* U$* F,-//_/I* 2$* %2;C"* 0$2'=C($V
3"#(2(%* =4* 4$)2'()4* B'1=714CD* B'(* !;%')$$3*
F(%(1)#'*:$;4C12=$4* =%* 2'14N(C*5$)* 1* %#'$71)V
%'=&* 2$* #1))"* $;)* %2;C=(%* 2$X1)C%* 1* 6'0D* f)=#*
!#O(4b=(* =%* 2'14N(C* 5$)* =3&)$@=48* 2'(* C)152*
314;%#)=&2D*
References
<1*W<*a14*C()D*/RS+*.*K2;C=(%*=4*Phyllosticta
/D*K2;C=(%*=4*!"#$7$8"*-9*/*.*//AD*
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F(2'=413* 69* K=@1)1314* OD* ,AAS* .* U$4=*
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5);=2* .* <* '$7=%2=#* )(@=(XD* J42()412=$417*
M$;)417*U$4=*F(%(1)#'*,9*/.+SD*
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L!9* C(* e);"2()* M9* !(55()2* M69* e)$(4(V
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citriasiana*%&D*4$@D9*2'(*#1;%(*$5*#=2);%*
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*
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**
++,*
Guignardia bispora and G. ellipsoidea spp. nov. and other Guignardia
species from palms (Arecaceae)
Wulandari NF1,2, To–Anun C.1*, McKenzie EHC3 and Hyde KD4,5
*
+-(&.)/0(1/*$2*34.1/*3./'$4$5"6*7.#84/"*$2*95):#84/8)(6*;':.15*!.:*<1:=()%:/"6*;':.15*!.:6*,+>??6*@'.:4.1AB*
>!:#)$C:$4$5"* -:=:%:$16* D(%(.)#'* ;(1/)(* 2$)* E:$4$5"6* F1A$1(%:.1* F1%/:/8/(* $2* G#:(1#(%6* ;:C:1$15* G#:(1#(* ;(1/)(* H4B*
D.".*H.I.)/.*E$5$)*J!B*KL6*;:C:1$15*+LM++6*F1A$1(%:.B*
N!.1..I:*O'(18.*P.1A#.)(*D(%(.)#'6*3):=./(*E.5*M>+Q?6*98#I4.1A6*R(S*T(.4.1AB*
KG#'$$4*$2*G#:(1#(6*!.(*7.'*P8.15*<1:=()%:/"6*NNN*!B*+B*@B*@.%8A*!8.15*-:%/):#/6*;':.15*D.:*,Q+??6*@'.:4.1AB*
,E$/.1"*.1A*!:#)$C:$4$5"*-(&.)/0(1/6*;$44(5(*$2*G#:(1#(6*J:15*G.8A*<1:=()%:/"6*D:".A'6*G.8A:*9).C:.B**
*
O84.1A.):* R76* @$U9181* ;6* !#J(1V:(* WX;* .1A* X"A(* J-B* >?++* U* Guignardia bispora*.1A*GB
ellipsoidea*%&&B*1$=B*.1A*$/'()*Guignardia*%&(#:(%*2)$0*&.40%*Y9)(#.#(.(ZB*!"#$%&'()(*>Y>Z6*++,U
+>[B*
*
@S$* Guignardia* %&(#:(%* #$44(#/(A* :1* 1$)/'()1* @'.:4.1A* A:22()* 0$)&'$4$5:#.44"* 2)$0* &)(=:$8%4"*
I1$S1* Guignardia* %&(#:(%* )(#$)A(A* $1* &.40%B* Guignardia bispora %&B* 1$=B* :%* A:%/:158:%'(A* C"*
'.=:15* /S$* .%#$%&$)(* /"&(%* .1A* GB ellipsoidea*%&B*1$=B*:%*A:%/:158:%'(A*C"*'.=:15*)(A8#(A*
08#:4.5:1$8%*.&&(1A.5(%*#$0&.)(A*/$*/'(*'$4$/"&(*$2*GB candeloflamma6*.4%$*2$81A*$1*&.40%B*@'(*
1(S*%&(#:(%*.)(*A(%#):C(A*.1A*:448%/)./(A*.1A*#$0&.)(A*S:/'*%:0:4.)*/.\.B*
*
Key words U* 9%#$0"#(/(%* U* E$/)"$%&'.():.#(.(* U* -$/':A(.4(%* U* P(.2* %&$/* U* 3./'$5(1* U*
@.\$1$0"*
*
Article Information**
D(#(:=(A*>>*-(#(0C()*>?+?*
9##(&/(A*K*7(C)8.)"*>?++*
38C4:%'(A*$14:1(*N+*!.)#'*>?++*
];$))(%&$1A:15*.8/'$)^*;'.:S./*@$_.181*U*(U0.:4*U*.5&&:??L`#':.150.:B.#B/'*
*
Introduction
Guignardia* %&(#:(%* $1* &.40%* '.=(* C((1*
)(4./:=(4"* S(44* %/8A:(A* YD('0* +M+K6* 9)\* a*
!b44()* +M,K6* 9.* +MQN6* 381:/'.4:15.0* +MQK6*
G:=.1(%.1* +M[K6* X.14:1* +MM?6* X"A(* +MM,6*
7)c'4:#'* a* X"A(* >???6* X"A(* (/* .4B* >???6*
d.11.*(/*.4B*>??+6*9.*a*e.1(=*>??>6*@."4$)*a*
X"A(* >??N6* 3:1)8.1* (/* .4B* >??QZ* .1A* 08#'* $2*
/'(* A./.* S.%* %800.):V(A* C"* X"A(* Y+MM,ZB*
X"A(* Y+MM,Z* :1/)$A8#(A* /S$* 1(S* %&(#:(%6*
)(A(%#):C(A* .* 28)/'()* %:\* %&(#:(%* 2)$0* &.40%6*
.1A* &)$=:A(A* .* I("* /$* /'(* &.40* Guignardia*
%&(#:(%B* @."4$)* a* X"A(* Y>??NZ* 2$81A* /')((*
%&(#:(%*$2*Guignardia*$1*&.40%6*(.#'*'.=:15*.*
Phyllosticta* .1.0$)&'* .1A* .* Leptodothiorella*
%&()0./:.4*%/./(B*G$1/:)./*(/*.4B*Y+MMKZ*)(#$)A(A*
$1(*%&(#:(%*$2*Phyllosticta*$##8)):15*$1*Areca*
%&B*:1*@'.:4.1AB*
Phyllosticta*%&(#:(%*'.=(*C((1*)(#$)A(A*
$1*&.40%*.%*(1A$&'"/(%*YP80"$15*(/*.4B*>??MZB*
PB cocoicola*:%* .*#$00$1*%&(#:(%* 2$81A* .%*.1*
(1A$&'"/(6* %.&)$C(* .1A* .* &./'$5(1* .1A* '.%*
Guignardia cocoicola* .%* /'(* /(4($0$)&'*
Y381:/'.4:15.0* +MQK6* @."4$)* +MMM6* X"A(* a*
@."4$)*>??N6*P80"$15*(/*.4B*>??MZB**
@'(* $Cf(#/:=(* $2* /':%* )(%(.)#'* S.%* /$*
:1=(%/:5./(* Guignardia*%&(#:(%*$##8):15*$1*
&.40%* :1* 1$)/'()1* @'.:4.1AB* F1* /':%* &.&()* S(*
A(%#):C(* /S$* 1(S* %&(#:(%* C.%(A* $1* 0$)&'$_
4$5:#.4*#'.).#/()%B*
Methods
Specimens
@'(* '$4$/"&(* %&(#:0(1* $2* Guignardia
candeloflamma HB* 7)c'4B* a* JB-B* X"A(* S.%

++L*
C$))$S(A*2)$0* EDF3*.1A* :%* :448%/)./(A*'()(* .%*
.%#$%&$)(%* .)(* #$0&.).C4(* /$* $1(* $2* /'(* 1(S*
%&(#:(%* 2)$0* &.40%B* 7)(%'* #$44(#/:$1%* $2* /'(*
1(S* Guignardia* %&(#:(%* S()(* #$44(#/(A* 2)$0*
;':.15*!.:*.1A*;':.15*D.:B**
*
Morphology
!:#)$%#$&"* S.%* #.)):(A* $8/* 8%:15* %/.1_
A.)A* /(#'1:g8(%B* 9%#$0./.* S()(* (\.0:1(A*
S:/'*.1*h4"0&8%*GTK?*0:#)$%#$&(6*.1A*0:#)$_
#'.).#/()%*(\.0:1(A* 8%:15*.* R:I$1* [?:*0:#)$_
%#$&(* S:/'* @.)$%$2/* &)$5).0* 2$)* 0(.%8):15*
%&$)(%B* 9* #.0().* 48#:A.* .//.#'(A* /$* .1*
h4"0&8%* ;i* K+* 0:#)$%#$&(* S.%* 8%(A* 2$)*
&)(&.)./:$1* $2* 4:1(* A).S:15%B* G&(#:0(1%* 2$)*
0:#)$%#$&:#* $C%()=./:$1* S()(* &)(&.)(A* C"*
'.1A* %(#/:$1:15B* P.#/$&'(1$4* #$//$1* C48(* .1A*
4.#/$54"#()$4* %$48/:$1* S()(* 8%(A* .%* 0$81/:15*
0(A:.B* -(/.:4%* $2* /.\$1$0:#* 1$=(4/:(%* .)(*
A(&$%:/(A*:1* !"#$E.1I* YSSSB!"#$E.1IB$)56*
;)$8%*(/*.4B*>??KZB*
*
Results
@S$*1(S* Guignardia* %&(#:(%*S()(*:A(1_
/:2:(A* .1A* .)(* A(%#):C(A6* :448%/)./(A* .1A* #$0_
&.)(A* S:/'* )(4./(A* %&(#:(%* 2)$0* &.40%B* Guig_
nardia candeloflamma*:%*.4%$*:448%/)./(A*.%*:/*
'.%* #$0&.).C4(* .%#$%&$)(%* /$* /'(* /S$* 1(S*
Guignardia*%&(#:(%*A(%#):C(A*$1*&.40%B*
*
Taxonomy
Guignardia bispora RB7B* O84.1A.):* a* JB-B*
X"A(6*sp. nov.* 7:5%*+U>6*QU>N6*KKUKM*
!"#$E.1I*!E*,+M?MQ**
W/"0$4$5"*U*1.0(A*2$)* /'(* /S$*/"&(%*$2*
.%#$%&$)(%B*
Guignardia candeloflamma* %:0:4:%* %(A*
.%#$%&$).(* C:%&$).(6* (44:&%$:A(.(* =(4* #"4:1A):_
#(.(6*+?U+L*j*NU,*=(4*+NU+K*j*NUK*k0B*
9%%$#:./(A*S:/'*C4:5'/*./*/'(*.&(\*.1A*$1*
/'(* 4.0:1.* $2* /'(* 4(.=(%6* 4(%:$1%* &.4(* C)$S16*
S:/'* /':16* A.)I* C)$S1* C$)A()6* 1(#)$/:#* /:%%8(%*
#$1/.:1:15* 180()$8%* #$1%&:#8$8%* .%#$0./.*
Y7:5%*+6*>6*Q6*[ZB*9%#$0./.*M,U+?,*k0*A:.0(_
/()6*++,U+>,*k0* ':5'6* $1*/'(*8&&()* %8)2.#(* $2*
/'(* 4(.=(%6* C4.#I6* 54$C$%(* /$* %8C54$C$%(6*
:00()%(A* :1* &4.1/* /:%%8(%6* #$):.#($8%6* %$4:/.)"*
/$*#48%/()(A6*$%/:$4./(6*$%/:$4(%*.%* C4.#I* A$/%*:1*
/'(* #(1/)(B* 3():A:80* +?U>,* k0* S:A(6* #$0_
&$%(A* $2* /S$* /$* /')((* 4."()%* $2* #(44%6* textura
angularis*.1A*&:50(1/(A*$8/S.)A4"*.1A*.)$81A*
$%/:$4(6* &.4()* :1%:A(* Y7:5%* M6* +?6* >+ZB* 3%(8A$_
&.).&'"%(%* %'$)/* #'.:1%* $2* 28%:2$)0* /$* $=$:A*
#(44%B* 9%#:* NLUK[* j* ++U+>* k0* Y
x
*l* K>* j* +>*
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*
Guignardia
arengae
Rehm
G. bispora N.F.
Wulandari &
K.D. Hyde
G. calami
(Syd. & P.
Syd.) Arx &
E. Müller
G. candelo-
flamma J.
Fröhl. &
K.D. Hyde
G. cocoɺs
(Petch) K.D.
Hyde
G. cocogena
(Cooke)
Punith.
G. ellipsoi-
dea N.F.
Wulandari
& K.D.
Hyde
G. manokwaria
K.D. Hyde
G. migrans
(Rehm) K.D.
Hyde
G. ryukyu-
ensis I. Hino
&
Katumoto
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08#:4.5:1$8%*.&&(1A.5(%B*@'(%(*#'.).#/()%*#.1*
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*
Acknowledgements
@':%* %/8A"* S.%* 281A(A* :1* &.)/* C"*
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R:4.0* O84.1A.):* :%* 5)./(284* /$* /'(* o).A8./(*
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/':%*)(%(.)#'B*
*
References
9.*X9*e.1*A()B*+MQN*U*G/8A:(%*:1*Phyllosticta
+B*G/8A:(%*:1*!"#$4$5"*,6*+U++?B*
9.*X9* e.1*A()6*e.1(=* GB*>??>* U* 9*D(=:%:$1*
$2* @'(* G&(#:(%* -(%#):C(AB* F1^* Phyllo-
stictaB* ;(1/)..4C8)(.8* =$$)* G#':00(4_
#84/8)(%6*</)(#'6*@'(*R(/'()4.1A%B*+UKMB*
9)\* H9* =$16* !b44()* WB* +M,K* U* -:(* o.//815(1*
A()* 90()$%&$)(%* 3")(1$0"#(/(1B* E(:/)*
J)"&/$5.0(124$).*G#'S(:V*++6*+UKNKB*
;)$8%* 3O6* o.0%* O6* G/.4&()%* H96* D$C()/* e6*
G/(5('8:%* oB* >??K* U* !"#$E.1I^* .1*
$14:1(*:1:/:./:=(*/$* 4.81#'* 0"#$4$5"*:1/$*
/'(*>+%/*#(1/8)"B*G/8A:(%*:1*!"#$4$5"*,?6*
+MU>>B*
7)c'4:#'* H6* X"A(* J-B* +MM,* U* Guignardia
candeloflamma*%&B*1$=B*#.8%:15*4(.2*
%&$/%* $2* Pinanga*%&B*!"#$4$5:#.4*
D(%(.)#'*MM6*++?U++>B*
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chinensisB*7815.4*-:=()%:/"*L6*+LQU+[?B*

ORIGINAL ARTICLE
Phyllosticta ophiopogonis sp. nov. from Ophiopogon
japonicus (Liliaceae)
S. Wikee
a
, N.F. Wulandari
a,b,c
, E.H.C. McKenzie
d
, K.D. Hyde
a,
*
a
School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
b
Department of Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai 51200, Thailand
c
Microbiology Division, Research Centre for Biology, Indonesian Institute of Sciences, Cibinong Science Centre,
Jl. Raya Jakarta Bogor KM. 46, Cibinong 16911, Indonesia
d
Manaaki Whenua Landcare Research, Private Bag 92170, Auckland, New Zealand
Received 9 July 2011; revised 8 October 2011; accepted 8 October 2011
Available online 19 October 2011
KEYWORDS
Coelomycetes;
Botryosphaeriaceae;
Dothideales;
Leaf spot;
Pathogen;
Taxonomy
Abstract A leaf spotting disease of an ornamental variety of Ophiopogon japonicus was discovered
at several locations in northern Thailand. In all cases a species of Phyllosticta was associated with
the lesions. Phyllosticta ophiopogonis sp. nov. is distinguished from Phyllosticta species from Lilia-
ceae in conidia size, mucilaginous sheath and appendage thus the species is introduced as new in this
paper. The new species which causes unsightly lesions on this ornamental plant is described, illus-
trated and compared with other similar Phyllosticta species.
ª2011 King Saud University. Production and hosting by Elsevier B.V. All rights reserved.
1. Introduction
The genus Phyllosticta had been relatively well-studied world-
wide and a monograph has been published with details of the
excepted species (van der Aa and Vanev, 2002). Five species
have been introduced since the publication of van der Aa
and Vanev (2002), mainly based on morphology and host
occurrence. Four new species were introduced from Japan by
Motohashi et al. (2008), while Phyllosticta citriasiana
Wulandari, Crous and Gruyter, from the peel of the fruit
Citrus maxima was introduced from China and Thailand
(Wulandari et al., 2009). Three new species of the sexual state
of Phyllosticta have also recently been introduced and include
Guignardia musicola, Wulandari, Cai and Hyde, Guignardia
bispora, Wulandari and Hyde, and Guignardia ellipsoidea,
Wulandari and Hyde from the palms in northern Thailand
(Wulandari et al., 2010a,b, 2011). Glienke et al. (2011) also
introduced Phyllosticta bifrenariae Pereira, Glienke and Crous
on orchids, Phyllosticta citribraziliensis Glienke and Crous on
Citrus and Phyllosticta brazilianiae, Stringri, Glienke and
Crous on Mangifera indica and epitypified Phyllosticta capital-
ensis Henn. and Phyllosticta citricarpa (McAlpine) Aa.
Ophiopogon japonicus (L.f.) Ker Gawl is an ornamental
plant grown in gardens and parks throughout northern
Thailand. During field surveys we repeatedly came across se-
verely diseased plants, with symptoms ranging from numerous
*Corresponding author. Tel.: +66 871791761.
E-mail address: kdhyde3@gmail.com (K.D. Hyde).
1319-562X ª2011 King Saud University. Production and hosting by
Elsevier B.V. All rights reserved.
Peer review under responsibility of King Saud University.
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King Saud University
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leaf spots to severe blighting (Fig. 1). The separation of
Phyllosticta species is presently mainly based on morphologi-
cal characters, although molecular data has recently helped
to differentiate some taxa, e.g. P. bifrenariae,P. brazilianiae,
P. citriasiana,P. citribraziliensis (Wulandari et al., 2009;
Glienke et al., 2011). However, despite the use of molecular
data, we still need to rely on morphological data (Hyde
et al., 2010; Udayanga et al., 2011).
There is no report of Phyllosticta species on Ophiopogon sp.
in northern Thailand or worldwide. We therefore compare the
species from O. japonicus with the species from other members
from Liliaceae. Phyllosticta ophiopogonis is however distinct.
The aim of this paper is therefore to introduce the new species
from O. japonicus based on morphology as it causes unsightly
disease of this ornamental. The new species is compared with
other Phyllosticta species described from Liliaceae in Table 1.
2. Material and Methods
Diseased leaves of O. japonicus were collected from various
sites in Chiang Rai Province, Thailand. Morphological charac-
ters were recorded using the methods described by Wulandari
et al. (2010a,b). Single spore isolates were prepared using the
method of Choi et al. (1999).
3. Results
Phyllosticta ophiopogonis Wulandari, Wikee and Hyde, sp.
nov. MycoBank: MB 519321 (Figs. 1A–N and 2M–O).
Etymology named after its host plant, O. japonicus.
Leaf spots on lamina and apex of the leaf, at first minute, later
becoming large spots with a dark reddish border, eventually
coalescing to form blights on the leaves, centre of lesions pale
Figure 1 (A–N) Phyllosticta ophiopogonis (MFLU11 0027, holotype). (A) Symptom of disease. (B–D) Pycnidia growing on infected leaf
of Liliaceae. (E) Conidiogenous cell. (F–J) Conidia Cross section through (K) Leptodothiorella state; scale bar = 50 lm. (L and M)
Spermatia; scale bar = 10 lm. (N) Upper of cultures after 5 weeks.
14 S. Wikee et al.

brown with numerous pycnidia (Fig. 1A–C). Pycnidia 96–99 lm
diameter, 75–81 lm high, on the surface of leaves, black, globose
to pyriform, immersed in plant tissues, coriaceous, solitary to
clustered, ostiolate, ostioles as black dots in the centre
(Fig. 1D). Conidiogenous cells 7–12 ·2–4 lmð!
x¼9#3lm;
n¼20Þ, holoblastic, determinate, discrete, hyaline, sometimes
rarely integrated, with cylindrical to doliiform cells lining the
pycnidial locule (Fig. 1E). Conidia 10–14 ·7–8 lmð!
x¼12 #7
lm;n¼20Þ, hyaline, 1-celled, coarse-guttulate, smooth-walled,
globose, ellipsoidal, clavate or obclavate, with an obtuse apex,
sometimes truncate at the base, surrounded by 0.8–1 lm thick
mucilaginous sheath which persists at maturity and with 5–
16 lm a single, hyaline, curved or straight appendage (Fig. 1H–J).
Leptodothiorella state, 60–80 lm in length, 40–50 lm in
wide and thick 22 lm. Spermatia are produced from spermati-
ogenous cells, cylindrical and globose at two ends 6–8 lm long,
6.7–8.3 ·1.3–1.6 lm.
Colonies black, fimbriate, black in reverse, reaching 3–5 cm
in diameter after 21 days incubation at 28 !C of on half
strength PDA.
Habitat: On living leaves causing leaf spots.
Host: O. japonicus (Lilliaceae).
Known distribution: Thailand (Chiang Rai).
Material examined: Thailand, Chiang Rai Province, Khun.
Korn Waterfall, on the leaves of O. japonicus, 10 November
2010, Wikee, WK 10 (MFLU11-0027, holotype); culture
ex-type MFLUCC11-0057; Wieng Chiang Rung, Houi Mae
Sak Waterfall, on the leaves of O. japonicus, 10 September
2010, S. Wikee, WK 12 (MFLU11-0028), culture
MFLUCC11-0059; Nang Lae, Pasangwiwat, on the leaves of
O. japonicus, 10 December 2010, Wikee, WK 17 (MFLU11-
0029), culture MFLUCC11-0063; ibid. 11 November 2010,
Wikee, WK 23 (MFLU11-0030), culture MFLUCC11-0069;
Weing Kan, on the leaves of O. japonicus, 6 January 2011,
Wikee, WK 26 (MFLU11-0031), culture MFLUCC10-0132.
Mae Fah Luang University, on the leaves of O. japonicus, 30
June 2010, Wulandari, NFW 330 (MFLU10-0480); ibid., 06
August 2010, Wulandari, NFW 332 (MFLU10-0482); ibid.,
Hue Pui Temple, on the leaves of O. japonicus, 28 October
2010, Wulandari, NFW 341 (MFLU10-0980); ibid., Hue
Pui Temple, on the leaves of O. japonicus, 17 August 2010,
Wulandari, NFW 343 (MFLU10-0982).
Notes: Van der Aa (1973) and van der Aa and Vanev
(2002) distinguished nine species of Phyllosticta on Liliaceae
(Table 1).
Of the Phyllosticta spp. that occurs on Liliaceae, P. cruenta
var. discincta is the most similar. The conidia of P. ophiopogonis
differ as they are smaller than P. cruenta var. discincta
(10–14 ·7–8 lmð!
x¼12 #7lm;n¼20Þversus 12–23 ·6.4–
10 lm) and the pycnidia are also smaller in P. ophiopogonis
(96–99 lm diameter, 75–81 lm high versus 100–195 lm diame-
ter 145 lm high). P. ophiopogonis also differs from Phyllosticta
hypoglossi in having shorter appendages, 5–16 lm versus 10 up
to 35 lm long (van der Aa, 1973).
Table 1 Phyllosticta spp. described from Liliaceae.
Phyllosticta
species
Host plant
and family
*
Pycnidia size
(lm)
Peridium
thickness
(lm)
Conidiogenous
cells (lm)
Conidia
(lm)
Sheath
size (lm)
Appendage
size (lm)
Reference
P. aspidistricola Aspidistra elatior var.
elatior Liliaceae
61–118 ·
86–110
– 7–12.5 ·
1.2–2.5
9.5–12.5 ·
8.5–10
– 17–24.5 Motohashi
et al.
(2008)
P. cruenta var.
discincta
Polygonatum latifolium,
Liliaceae
100–195 ·
145
9–21 4–14 ·3–6 12–23 ·
6.4–10
0.2 4–17 Bissett
(1979a)
present
study
P. crypta Smilax sp.,
Liliaceae
70–130 ·
45–95
4–14 5–12 ·
2–3.5
5.4–8.9 ·
3.8–6.2
0.3–1.0 3–8 Bissett
(1979b)
P. cumminsii Smilax sp.,
Liliaceae
75–140 ·
110
4–19 3.5–14 ·3–6 6.7–10.5 1–2 5–20 Bissett
(1979b)
P. discincta Uvularia grandiflora,
Liliaceae
65–120 5–12 4–13 ·
2.5–6
5–8.6 ·
3.9–6.6
Less than 0.8 4–14 van der
Aa and
Vanev
(2002)
P. hemerocallidis Hemerocallis fulva,
Liliaceae
84–139 – – 8–13 ·3–5 – 3–10 van der
Aa and
Vanev
(2002)
P. hypoglossi Ruscus hypoglossum,
Liliaceae
120–250 12–30 4–10 ·
2–3.5
8–15 (%18) ·
6–10
– 10 up to 35 Van der
Aa
(1973)
P. ophiopogonis Ophiopogon japonicus,
Liliaceae
96–100 ·
75–80
9–15 7–12 ·2–4 10–14 ·7–8 0.8–1 5–16 Present
study
P. subeffusa Smilax herbacca,
Liliaceae
90–150 ·
120–140
5–16 5–8 ·3–4.5 7–13 ·7–10 – 5–7 up to 15 Van der
Aa (1973)
P. yuccae Yucca elephantipes,
Liliaceae
90–150 14–38 5.4–9.8 ·
2.7–6
7.5–15.4 ·6–9.5 1 4–15 Bissett
(1986)
*
They may now belong to other families.
Phyllosticta ophiopogonis sp. nov. from Ophiopogon japonicus (Liliaceae) 15

4. Discussion
There is a move towards use of one name for a single biological
species instead of different names for different morphs (Hyde
et al., 2011). In this study, the sexual Guignardia state did
not form in any of the collections made or in culture. We
choose to use the oldest name Phyllosticta as compared to
Guignardia for this taxon as Phyllosticta species usually cause
serious disease. If the teleomorph is found later it can be de-
scribed under P. ophiopogonis. This disease is important as it
causes unsightly spots and blights on this commonly used or-
namental (Fig. 1A).
Acknowledgements
The Global Research Network for Fungal Biology and King
Saud University are thanked for supporting this research.
Nilam Wulandari is grateful to the Mushroom Research Foun-
dation for a scholarship to carry our studies towards a Ph.D.
and the Graduate School, Chiang Mai University, Thailand
for the financial support and the School of Science, Mae Fah
Luang University for the laboratory facilities. MFLU awarded
Grant No. 53101020017 to study the genus Phyllosticta in
northern Thailand and the National Research Council of
Thailand awarded Grant No. 54201020004 to study the genus
Phyllosticta in Thailand. The Thailand Research Fund in the
Royal Golden Ph.D. Jubilee Program agreement No. Ph.D. /
0198/2552 in 2.B.M.F./52/A.1.N.XX to study the taxonomy
and phylogeny of Phyllosticta is acknowledged. Crous, CBS,
the Netherlands is thanked for partially funding this research.
Wara Asfiya (LIPI) and Samantha Karunarathna (MFLU) are
thanked for valuable references on Phyllosticta species.
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Figure 2 (M–O) Phyllosticta ophiopogonis (MFLU 10-0480) line
drawing. (M) Section of pycnidia in the leaf (darkened area are
plant cells – arrowed). (N) Conidiogenous cells. (O) Conidia with
appendage and sheath.
16 S. Wikee et al.

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