diaporthe: a genus of endophytic, saprobic and plant ...2 persoonia–volume31,2013 diaporthe...
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Persoonia 31, 2013: 1–41www.ingentaconnect.com/content/nhn/pimj http://dx.doi.org/10.3767/003158513X666844RESEARCH ARTICLE
INTRODUCTION
Species of Diaporthe and their Phomopsis asexual states have broad host ranges and are widely distributed, occurring as plant pathogens, endophytes or saprobes, but also as pathogens of humans and other mammals (Webber&Gibbs1984,Carroll1986,Boddy&Griffith1989,Rehner&Uecker1994,Garcia-Reyneetal.2011,Udayangaetal.2011).Diaporthespp.areresponsible for diseases on a wide range of plants hosts, some of which are economically important worldwide, causing root and fruit rots, dieback, cankers, leaf spots, blights, decay and wilt(Uecker1988,Mostertetal.2001a,vanRensburgetal.2006,Santosetal.2011,Thompsonetal.2011).Currently,MycoBank (accessedSept.2012) listsmore than1 000 names in the genus Phomopsis, while Diaporthe contains morethan860names.Inthepastspecieshavechieflybeendescribedundertheassumptiontheyarehost-specific,leadingto a proliferation of names based on the hosts from which they were isolated (Uecker 1988).However, subsequent studieshave found that many species are able to colonise diverse hosts as opportunists, and that several different species could even co-occuronthesamehostorlesion(Brayford1990,Rehner&Uecker1994,Mostertetal.2001a,Farretal.2002,Crous&Groenewald2005).Curiously,somespeciesofDiaporthe can be either pathogenic or harmless endophytes depending on the
hostanditshealth.Forexample,D. phaseolorum is pathogenic tosoybean(Santosetal.2011),butendophyticinmangroves(Laguncularia racemosa) (Sebastianeet al. 2011).With thedeletionofArt.59fromtheInternationalCodeofNomenclatureforalgae,fungi,andplants(ICN),asexualandsexualnamesoffungireceiveequalstatus(Hawksworthetal.2011,Wingfieldet al. 2012).Because the nameDiaporthe (1870) predatesPhomopsis(1905),Diaporthe is adopted in the present study forthisgroupoffungi(Santosetal.2010,2011,Crousetal.2011,Udayangaetal.2012).Diaporthe (incl. itsPhomopsis state) has been reported asoneofthemostfrequentlyencounteredgeneraofendophyticfungiinseveralplanthosts(Muralietal.2006,Botella&Diez2011).Thegenushasalsofrequentlybeenrecognisedasaproducerof interestingenzymesandsecondarymetabolites(Isaka et al. 2001,Kobayashi et al. 2003,Dai et al. 2005,Elsaesseretal.2005)withantibiotic(Bandre&Sasek1977,Dettrakuletal.2003,Linetal.2005)oranticancer(Kumaran&Hur2009)activity.Furthermore,speciesofDiaporthe have in thepastbeennotedtodeterherbivory(Brayford1990,Weber2009,Vesterlundetal.2011),havelignocellulolyticactivities(Jordaanetal.2006),orhavebeenappliedasbioherbicides(Ashetal.2010).Theaccurateapplicationofacceptednamesofplantpathogenicfungi is essential for the development of effective biosecurity and tradepolicies (Crous&Groenewald2005,Wingfieldetal.2012).Thetaxonomyofmanygroupsofplantpathogenicfungihasinthepastbeenbasedonhostassociation(Crousetal.2013,Groenewaldetal.2013).AlthoughsomespeciesofDiaporthearehostspecific,agreatnumberhavebeennotedtooccuronmorethanonehost(Brayford1990,Rehner&Uecker1994,Farretal.2002).Similarobservations ledWehmeyer
Diaporthe: a genus of endophytic, saprobic and plant pathogenic fungiR.R.Gomes1,C.Glienke1,S.I.R.Videira2,L.Lombard2,J.Z.Groenewald 2, P.W.Crous 2,3,4
1 DepartmentofGenetics,UniversidadeFederaldoParaná,CentroPolitéc-nico,Box19071,81531-990,Curitiba,Brazil.
2 CBS-KNAWFungalBiodiversityCentre,Uppsalalaan8,3584CT,Utrecht,TheNetherlands;correspondingauthore-mail:[email protected].
3Microbiology,DepartmentofBiology,UtrechtUniversity,Padualaan8,3584CHUtrecht,TheNetherlands.
4WageningenUniversityandResearchCentre(WUR),LaboratoryofPhyto-pathology,Droevendaalsesteeg1,6708PBWageningen,TheNetherlands.
Key words
DiaporthalesDiaportheMulti-LocusSequenceTyping(MLST)Phomopsissystematics
Abstract Diaporthe(Phomopsis)specieshaveoftenbeenreportedasplantpathogens,non-pathogenicendo-phytesorsaprobes,commonlyisolatedfromawiderangeofhosts.Theprimaryaimofthepresentstudywastoresolve the taxonomy and phylogeny of a large collection of Diaporthe species occurring on diverse hosts, either aspathogens,saprobes,orasharmlessendophytes.Inthepresentstudyweinvestigated243isolatesusingmulti-locusDNAsequencedata.AnalysesoftherDNAinternaltranscribedspacer(ITS1,5.8S,ITS2)region,andpartialtranslationelongationfactor1-alpha(TEF1),beta-tubulin(TUB),histoneH3(HIS)andcalmodulin(CAL)genesresolved95clades.Fifteennewspeciesaredescribed,namelyDiaporthe arengae, D. brasiliensis, D. endophytica, D. hongkongensis, D. inconspicua, D. infecunda, D. mayteni, D. neoarctii, D. oxe, D. paranensis, D. pseudomangiferae, D. pseudophoenicicola, D. raonikayaporum, D. schini and D. terebinthifolii.Afurther14newcombinationsare introduced in Diaporthe, and D. anacardiiisepitypified.AlthoughspeciesofDiaporthe have in the past chiefly beendistinguishedbasedonhostassociation,resultsofthisstudyconfirmseveraltaxatohavewidehostranges,suggestingthattheymovefreelyamonghosts,frequentlyco-colonisingdiseasedordeadtissue.Incontrast,someplantpathogenicandendophytictaxaappeartobestrictlyhostspecific.Giventhisdiverseecologicalbehaviouramong members of Diaporthe,futurespeciesdescriptionslackingmoleculardata(atleastITSandHISorTUB)shouldbestronglydiscouraged.
Article infoReceived:27November2012;Accepted:24February2013;Published:28March2013.
2 Persoonia–Volume31,2013
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Tabl
e 1H
ost/substrate,locality,collectorandGenBankaccessionnumbersofstrainsincludedinthestudy.
Spe
cies
O
rigin
al n
ame
Stra
in1
Isolationsource
Hostfam
ily
Locality
Collector
GenBa
nkAccessionnum
bers(ITS
,CAL
,HIS,TEF
1,TUB)
2
3R.R.Gomesetal.:Diaporthe
Dia
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d Fa
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KC343295
KC343537
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Dia
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P. c
onvo
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iCBS124654;DP0727
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volv
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KC343296
KC343538
KC343780KC344022
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rata
egi
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L.H
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KC343539
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eber
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KC343298
KC343540
KC343782KC344024
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-U5431;
Asp
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Faba
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J.C.JansevanRensburgKC343057
KC343299
KC343541
KC343783KC344025
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tea
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rote
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arincowitz
KC343058
KC343300
KC343542
KC343784KC344026
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eced
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llana
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oryl
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KC343059
KC343301
KC343543
KC343785KC344027
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. dec
eden
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ave
llana
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oryl
acea
eSweden
K.&
L.H
olm
KC343060
KC343302
KC343544
KC343786KC344028
Dia
porth
e de
trusa
D
. det
rusa
CBS109770;AR3424
Ber
beris
vul
garis
B
erbe
ridac
eae
Austria
A.Y.R
ossm
an
KC343061
KC343303
KC343545
KC343787KC344029
D
. det
rusa
CBS114652;UPSC3371
Ber
beris
vul
garis
B
erbe
ridac
eae
Sweden
K.&
L.H
olm
KC343062
KC343304
KC343546
KC343788KC344030
D
. det
rusa
CBS140.27
Ber
beris
vul
garis
B
erbe
ridac
eae
–L.E.W
ehmeyer
KC343063
KC343305
KC343547
KC343789KC344031
Dia
porth
e el
aeag
ni
P. e
laea
gni
CBS504.72
Ela
eagn
ussp.,twig
Ela
eagn
acea
eNetherlands
J.Gremmen
KC343064
KC343306
KC343548
KC343790KC344032
Dia
porth
e en
doph
ytic
a,sp.nov.
–CBS133811;LGMF916;
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343065
KC343307
KC343549
KC343791KC344033
CPC20292(ex-type)
–
LGMF911;C
PC20287
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343066
KC343308
KC343550
KC343792KC344034
–
LGMF919;C
PC20295
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343067
KC343309
KC343551
KC343793KC344035
–
LGMF928;C
PC20304
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343068
KC343310
KC343552
KC343794KC344036
–
LGMF934;C
PC20310
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343069
KC343311
KC343553
KC343795KC344037
–
LGMF935;C
PC20311
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343070
KC343312
KC343554
KC343796KC344038
–
LGMF937;C
PC20313
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343071
KC343313
KC343555
KC343797KC344039
–
LGMF948;C
PC20324
Gly
cine
max
, see
d Fa
bace
ae
Brazil
A.Almeida
KC343072
KC343314
KC343556
KC343798KC344040
Dia
porth
e er
es
D. e
res
CBS101742
Frax
inussp.,fallenfruit
Ole
acea
eNetherlands
G.J.M.Verkley
KC343073
KC343315
KC343557
KC343799KC344041
D
. med
usae
aCBS102.81
Jugl
ans
regi
a, tw
ig
Jugl
anda
ceae
Italy
M.B
isiach
KC343074
KC343316
KC343558
KC343800KC344042
D
. ere
sCBS109767;AR3538;W
J1643
Ace
r cam
pest
re
Ace
race
ae
Austria
W.Jaklitsch
KC343075
KC343317
KC343559
KC343801KC344043
D
. arc
tii
CBS110.85
Arc
tiumsp.,deadstem
sA
ster
acea
eNetherlands
M.deNooij
KC343076
KC343318
KC343560
KC343802KC344044
P.
ski
mm
iae
CBS122.82
Ski
mm
ia ja
poni
ca, d
ying
twig
s R
utac
eae
Netherlands
H.A.v.K
esteren
KC343077
KC343319
KC343561
KC343803KC344045
P
hom
opsi
ssp.no.23CBS129168
Rho
dode
ndro
nsp.
Eric
acea
eLatvia
I.Apine
KC343078
KC343320
KC343562
KC343804KC344046
D
. con
orum
CBS186.37
Pic
ea a
bies
, see
dlin
g P
inac
eae
UK
T.R.P
eace
KC343079
KC343321
KC343563
KC343805KC344047
P.
con
trove
rsa
CBS250.38
Frax
inus
exc
elsi
or, l
ivin
g an
d de
ad tw
ig
Ole
acea
eUK:S
cotland
J.A.M
acDonald
KC343080
KC343322
KC343564
KC343806KC344048
P.
stic
tica
CBS267.32
––
–W.G.H
utchinson
KC343081
KC343323
KC343565
KC343807KC344049
P.
rudi
sCBS267.55
Labu
rnum
× w
ater
eri ‘
Voss
ii’
Faba
ceae
Netherlands
I.deBoer
KC343082
KC343324
KC343566
KC343808KC344050
P.
rano
jevi
cii
CBS283.85
Alli
um g
igan
teum
, dea
d st
em
Alli
acea
eNetherlands
H.A.vanderAa
KC343083
KC343325
KC343567
KC343809KC344051
D
. ere
sCBS287.74
Sor
bus
aucu
paria
, dea
d br
anch
R
osac
eae
Netherlands
W.M.Loerakker
KC343084
KC343326
KC343568
KC343810KC344052
P.
osm
anth
iCBS297.77
Osm
anth
us a
quifo
lium
, lea
f tip
O
leac
eae
Netherlands
H.A.vanderAa
KC343085
KC343327
KC343569
KC343811
KC344053
P.
cac
tiCBS365.97
Opu
ntiasp.,cladodes
Cac
tace
ae
Netherlands
H.A.vanderAa
KC343086
KC343328
KC343570
KC343812KC344054
P.
cru
stos
aCBS370.67;MUCL9931
Ilex
aqui
foliu
m, d
ead
leaf
A
quifo
liace
ae
Netherlands
H.A.vanderAa
KC343087
KC343329
KC343571
KC343813KC344055
D
. per
nici
osa
CBS375.61
Mal
us s
ylve
stris
, rot
ten
fruit
Ros
acea
e–
Geigy
KC343088
KC343330
KC343572
KC343814KC344056
P.
pha
seol
iCBS422.50
Pha
seol
us v
ulga
ris
Faba
ceae
Netherlands
Goossens
KC343089
KC343331
KC343573
KC343815KC344057
P.
cot
onea
stri
CBS439.82;BBAP-407;IMI162181a
Cot
onea
stersp.
Ros
acea
eUK:S
cotland
H.B
utin
KC343090
KC343332
KC343574
KC343816KC344058
(isotypeofP
hom
opsi
s co
tone
astri)
P.
cru
cife
rae
CBS445.62
Alli
aria
offi
cina
lis
Bra
ssic
acea
eNetherlands
G.H.B
oerema
KC343091
KC343333
KC343575
KC343817KC344059
P.
dur
andi
ana
CBS485.96
Rum
ex h
ydro
lapa
thum
, dea
d st
em
Pol
ygon
acea
eNetherlands
H.A.vanderAa
KC343092
KC343334
KC343576
KC343818KC344060
D
. sep
osita
CBS528.83
Wis
teria
sin
ensi
s, d
ead
bran
ch
Faba
ceae
Netherlands
H.A.vanderAa
KC343093
KC343335
KC343577
KC343819KC344061
P.
abu
tilon
is
CBS688.97
Abu
tilonsp.
Mal
vace
ae
Netherlands
A.Aptroot
KC343094
KC343336
KC343578
KC343820KC344062
P.
cru
stos
aCBS694.94
Ilex
aqui
foliu
m, t
wig
s su
fferin
g fro
m d
ieba
ck A
quifo
liace
ae
Netherlands
G.J.M.Verkley
KC343095
KC343337
KC343579
KC343821KC344063
P.
mag
nolii
cola
CBS791.68
Mag
nolia
× s
oula
ngea
na, w
ither
ing
leaf
M
agno
liace
ae
Netherlands
H.A.vanderAa
KC343096
KC343338
KC343580
KC343822KC344064
P.
triti
ci
CBS841.84
Hor
deum
sp.,leafspot
Poa
ceae
Germany
M.H
ossfeld
KC343097
KC343339
KC343581
KC343823KC344065
Dia
porth
e eu
geni
ae,com
b.nov.
P. e
ugen
iae
CBS444.82
Eug
enia
aro
mat
ica,
leaf
M
yrta
ceae
WestS
umatra
R.K
asim
KC343098
KC343340
KC343582
KC343824KC344066
Dia
porth
e fib
rosa
D
. fib
rosa
CBS109751;AR3425
Rha
mnu
s ca
thar
tica
Rha
mna
ceae
Austria
A.Y.R
ossm
an
KC343099
KC343341
KC343583
KC343825KC344067
D
. fib
rosa
CBS113830;UPSC2117
Rha
mnu
s ca
thar
tica
Rha
mna
ceae
Sweden
K.&
L.H
olm
KC343100
KC343342
KC343584
KC343826KC344068
Dia
porth
e fo
enic
ulac
ea
D. f
oeni
cula
cea
CBS111553
Foen
icul
um v
ulga
re, b
ase
of s
enes
cent
A
piac
eae
Spain
A.J.L.P
hillips
KC343101
KC343343
KC343585
KC343827KC344069
st
em
D. f
oeni
cula
cea
CBS111554
Foen
icul
um v
ulga
re, b
ase
of s
enes
cent
A
piac
eae
Portugal
A.J.L.P
hillips
KC343102
KC343344
KC343586
KC343828KC344070
st
em
P. th
eico
la
CBS116957;NZ-37
Pyr
us p
yrifo
lia
Ros
acea
eNew
Zealand
W.K
andula
KC343103
KC343345
KC343587
KC343829KC344071
D
. neo
thei
cola
CBS123208;Di-C
004/5(ex-type
Foen
icul
um v
ulga
re
Api
acea
ePortugal
A.J.L.P
hillips
KC343104
KC343346
KC343588
KC343830KC344072
of D
. neo
thei
cola)
D
. neo
thei
cola
CBS123209;Di-C
004/4(ex-typeof
Foen
icul
um v
ulga
re
Api
acea
ePortugal
A.J.L.P
hillips
KC343105
KC343347
KC343589
KC343831KC344073
D. n
eoth
eico
la)
P.
mal
if.sp.a
myg
dali
CBS171.78
Pru
nus
amyg
dalu
s, d
ried
fruit
Ros
acea
eItaly
A.C
iccarone
KC343106
KC343348
KC343590
KC343832KC344074
4 Persoonia–Volume31,2013
P.
thei
cola
CBS187.27(ex-typeofP
. the
icol
a)
Cam
ellia
sin
ensi
s, le
aves
and
bra
nche
s Th
eace
ae
Italy
M.C
urzi
KC343107
KC343349
KC343591
KC343833KC344075
P.
dio
spyr
iCBS287.56
Dio
spyr
os k
aki,
twig
, afte
r fro
st d
amag
e E
bena
ceae
Italy
M.R
ibaldi
KC343108
KC343350
KC343592
KC343834KC344076
D
. sep
osita
CBS357.69
Wis
teria
sin
ensi
s, d
ead
twig
s Fa
bace
ae
Netherlands
H.A.vanderAa
KC343109
KC343351
KC343593
KC343835KC344077
P.
cas
uarin
ae
CBS400.48
––
India
S.R.B
ose
KC343110
KC343352
KC343594
KC343836KC344078
P.
bou
gain
ville
ae
CBS603.88
Bou
gain
ville
a sp
ecta
bilis
, ped
uncl
es o
f N
ycta
gina
ceae
Portugal
H.A.vanderAa
KC343111
KC343353
KC343595
KC343837KC344079
flo
wer
s D
iapo
rthe
ganj
ae,com
b.nov.
P. g
anja
eCBS180.91;ILLS
43621(ex-type)
Can
nabi
s sa
tiva,
dea
d le
af
Can
naba
ceae
USA:Illinois
J.M.M
cPartland
KC343112
KC343354
KC343596
KC343838KC344080
Dia
porth
e ga
rden
iae,com
b.nov.
P. g
arde
niae
CBS288.56
Gar
deni
a flo
rida,
ste
m
Rub
iace
ae
Italy
M.R
ibaldi
KC343113
KC343355
KC343597
KC343839KC344081
Dia
porth
e he
liant
hi
D. h
elia
nthi
CBS344.94
Hel
iant
hus
annu
us, s
eed
Ast
erac
eae
––
KC343114
KC343356
KC343598
KC343840KC344082
D
. hel
iant
hi
CBS592.81(ex-type)
Hel
iant
hus
annu
us, o
verw
inte
ring
stem
A
ster
acea
eSerbia
M.M
untañola-CvetkovicKC343115
KC343357
KC343599
KC343841KC344083
Dia
porth
ecf.h
evea
e 1
P. h
evea
eCBS852.97
Hev
ea b
rasi
liens
is
Eup
horb
iace
ae
Brazil
D.S.Attili
KC343116
KC343358
KC343600
KC343842KC344084
Diaporthecf.h
evea
e 2
P. h
evea
eCBS681.84
Hev
ea b
rasi
liens
is, l
eaf
Eup
horb
iace
ae
India
K.Jayarathnam
KC343117
KC343359
KC343601
KC343843KC344085
Dia
porth
e hi
ckor
iae
D. h
icko
riae
CBS145.26(ex-type)
Car
ya g
labr
a Ju
glan
dace
ae
USA:M
ichigan
L.E.W
ehmeyer
KC343118
KC343360
KC343602
KC343844KC344086
Dia
porth
e ho
ngko
ngen
sis,sp.nov.
P. p
ittos
pori
CBS115448;HKUCC9104;
Dic
hroa
febr
ífuga
, fru
it H
ydra
ngea
ceae
HongKong
K.D.H
yde
KC343119
KC343361
KC343603
KC343845KC344087
AT646DF24(ex-type)
Dia
porth
e ho
rdei,com
b.nov.
P. h
orde
iCBS481.92
Hor
deum
vul
gare
, roo
t P
oace
ae
Norway
L.Sundheim
KC343120
KC343362
KC343604
KC343846KC344088
Dia
porth
e im
puls
a D
. im
puls
aCBS114434;UPSC3052
Sor
bus
aucu
paria
R
osac
eae
Sweden
K.&
L.H
olm
KC343121
KC343363
KC343605
KC343847KC344089
D
. im
puls
aCBS141.27
Sor
bus
amer
ican
a R
osac
eae
–L.E.W
ehmeyer
KC343122
KC343364
KC343606
KC343848KC344090
Dia
porth
e in
cons
picu
a,sp.nov.
Dia
porth
esp.
CBS133813;LGMF930;
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343123
KC343365
KC343607
KC343849KC344091
CPC20306(ex-type)
D
iapo
rthesp.
LGMF922;C
PC20298
Spo
ndia
s m
ombi
n A
naca
rdia
ceae
Brazil
K.R
odriguez
KC343124
KC343366
KC343608
KC343850KC344092
D
iapo
rthesp.
LGMF931;C
PC20307
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343125
KC343367
KC343609
KC343851KC344093
Dia
porth
e in
fecu
nda,sp.nov.
Dia
porth
esp.
CBS133812;LGMF906;
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343126
KC343368
KC343610
KC343852KC344094
CPC20282(ex-type)
D
iapo
rthesp.
LGMF908;C
PC20284
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343127
KC343369
KC343611
KC343853KC344095
D
iapo
rthesp.
LGMF912;C
PC20288
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343128
KC343370
KC343612
KC343854KC344096
D
iapo
rthesp.
LGMF917;C
PC20293
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343129
KC343371
KC343613
KC343855KC344097
D
iapo
rthesp.
LGMF918;C
PC20294
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343130
KC343372
KC343614
KC343856KC344098
D
iapo
rthesp.
LGMF920;C
PC20296
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343131
KC343373
KC343615
KC343857KC344099
D
iapo
rthesp.
LGMF933;C
PC20309
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343132
KC343374
KC343616
KC343858KC344100
D
iapo
rthesp.
LGMF940;C
PC20316
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343133
KC343375
KC343617
KC343859KC344101
Dia
porth
e ju
glan
dina
D
. jug
land
ina
CBS121004;DP0659
Jugl
anssp.,deadwood
Jugl
anda
ceae
USA:TennesseeL.Vasilyeva
KC343134
KC343376
KC343618
KC343860KC344102
Dia
porth
e lo
ngis
pora,com
b.nov.
D. s
trum
ellavar.
CBS194.36(ex-type)
Rib
essp.
Gro
ssul
aria
ceaeCanada
L.E.W
ehmeyer
KC343135
KC343377
KC343619
KC343861KC344103
lo
ngis
pora
D
iapo
rthe
lusi
tani
cae
D. l
usita
nica
eCBS123212;Di-C
001/5(ex-type)
Foen
icul
um v
ulga
re, s
tem
A
piac
eae
Portugal
J.M.S
antos
KC343136
KC343378
KC343620
KC343862KC344104
D
. lus
itani
cae
CBS123213;Di-C
001/3
Foen
icul
um v
ulga
re, s
tem
A
piac
eae
Portugal
J.M.S
antos
KC343137
KC343379
KC343621
KC343863KC344105
Dia
porth
e m
anih
otia
P.
man
ihot
CBS505.76
Man
ihot
util
issi
ma,
leav
es
Eup
horb
iace
ae
Rwanda
J.Sem
al
KC343138
KC343380
KC343622
KC343864KC344106
Dia
porth
e m
ayte
ni,sp.nov.
–CBS133185;LGMF938;
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343139
KC343381
KC343623
KC343865KC344107
CPC20314(ex-type)
Dia
porth
e m
egal
ospo
ra
D. m
egal
ospo
ra
CBS143.27
Sam
bucu
s ca
nade
nsis
C
aprif
olia
ceae
–L.E.W
ehmeyer
KC343140
KC343382
KC343624
KC343866KC344108
Dia
porth
e m
elon
is
D. p
hase
olor
um
CBS435.87
Gly
cine
soj
a Fa
bace
ae
Indonesia
H.V
ermeulen
KC343141
KC343383
KC343625
KC343867KC344109
var.
soja
e
D. m
elon
is
CBS507.78(ex-isotype)
Cuc
umis
mel
o C
ucur
bita
ceae
USA:Texas
L.Beraha&M.J.O
’BrienKC343142
KC343384
KC343626
KC343868KC344110
Dia
porth
e m
usig
ena
D. m
usig
ena
CBS129519;CPC17026(ex-type)
Mus
asp.,leaves
Mus
acea
eAustralia
P.W.C
rous&R.G.S
hivasKC343143
KC343385
KC343627
KC343869KC344111
Dia
porth
e ne
illia
e D
. nei
lliae
CBS144.27
Spi
raeasp.
Ros
acea
e–
L.E.W
ehmeyer
KC343144
KC343386
KC343628
KC343870KC344112
Dia
porth
e ne
oarc
tii,sp.nov.
D. a
rctii
CBS109490;GB6421;AR3450(ex-type)
Am
bros
ia tr
ifida
A
ster
acea
eUSA:
G.B
ills
KC343145
KC343387
KC343629
KC343871KC344113
New
Jersey
Dia
porth
e no
bilis
P.
cas
tane
aCBS113470;DAOM226800
Cas
tane
a sa
tiva,
che
stnu
ts c
olle
cted
in
Faga
ceae
Korea
K.A.S
eifert
KC343146
KC343388
KC343630
KC343872KC344114
gr
ocer
y st
ore
P.
fuku
shii
CBS116953;NZ-26
Pyr
us p
yrifo
lia
Ros
acea
eNew
Zealand
W.K
andula&
KC343147
KC343389
KC343631
KC343873KC344115
L.Castlebury
P.
fuku
shii
CBS116954;NZ-27
Pyr
us p
yrifo
lia
Ros
acea
eNew
Zealand
W.K
andula&
KC343148
KC343390
KC343632
KC343874KC344116
L.Castlebury
D
. per
nici
osa
CBS124030;GJS77-49
Mal
us p
umila
, bar
k R
osac
eae
New
Zealand
G.J.S
amuels
KC343149
KC343391
KC343633
KC343875KC344117
P
hom
opsi
ssp.no.22CBS129167
Rho
dode
ndro
nsp.
Eric
acea
eLatvia
I.Apine
KC343150
KC343392
KC343634
KC343876KC344118
D
. nob
ilis
CBS200.39
Laur
us n
obili
s, s
tem
La
urac
eae
Germany
Kotthoff
KC343151
KC343393
KC343635
KC343877KC344119
D
. pul
la
CBS338.89
Hed
era
helix
A
ralia
ceae
Yugoslavia
M.M
untañola-CvetkovicKC343152
KC343394
KC343636
KC343878KC344120
Tabl
e 1(cont.)
Spe
cies
O
rigin
al n
ame
Stra
in1
Isolationsource
Hostfam
ily
Locality
Collector
GenBa
nkAccessionnum
bers(ITS
,CAL
,HIS,TEF
1,TUB)
2
5R.R.Gomesetal.:Diaporthe
P. c
onor
um
CBS587.79
Pin
us p
enta
phyl
la,bonzaiimportedfrom
Pin
acea
eJapan
G.H.B
oerema
KC343153
KC343395
KC343637
KC343879KC344121
JapanintoNetherlands
Dia
porth
e no
mur
ai
D. n
omur
ai
CBS157.29
Mor
ussp.
Mor
acea
eJapan
K.Togashi
KC343154
KC343396
KC343638
KC343880KC344122
Dia
porth
e no
vem
D
. nov
em
CBS127269;5-27/3-1
Gly
cine
max
, see
d Fa
bace
ae
Croatia
T.Duvnjak
KC343155
KC343397
KC343639
KC343881KC344123
D
. nov
em
CBS127270;4-27/3-1(ex-type)
Gly
cine
max
, see
d Fa
bace
ae
Croatia
T.Duvnjak
KC343156
KC343398
KC343640
KC343882KC344124
D
. nov
em
CBS127271;5/27/3-3
Gly
cine
max
, see
d Fa
bace
ae
Croatia
T.Duvnjak
KC343157
KC343399
KC343641
KC343883KC344125
D
. par
dalo
ta
CBS354.71
Pol
ygon
atum
odo
ratu
m, l
eave
s C
onva
llaria
ceaeRom
ania
O.C
onstantinescu
KC343158
KC343400
KC343642
KC343884KC344126
D
iapo
rthesp.
LGMF943;C
PC20319
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343159
KC343401
KC343643
KC343885KC344127
Dia
porth
e on
cost
oma
D. o
ncos
tom
aCBS100454
Rob
inia
pse
udoa
caci
a, le
af s
pot
Faba
ceae
Germany
H.B
utin
KC343160
KC343402
KC343644
KC343886KC344128
D
. onc
osto
ma
CBS109741;AR3445
Rob
inia
pse
udoa
caci
a Fa
bace
ae
Russia
L.Vasilyeva
KC343161
KC343403
KC343645
KC343887KC344129
D
. onc
osto
ma
CBS589.78
Rob
inia
pse
udoa
caci
a, d
ead
bran
ches
Fa
bace
ae
France
H.A.vanderAa
KC343162
KC343404
KC343646
KC343888KC344130
P.
cru
stos
aCBS809.85
Ilex
aqui
foliu
m, l
eaf
Aqu
ifolia
ceae
Germany
M.H
esse
KC343163
KC343405
KC343647
KC343889KC344131
Dia
porth
e ox
e,sp.nov.
–CBS133186;LGMF942;
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343164
KC343406
KC343648
KC343890KC344132
CPC20318(ex-type)
–
CBS133187;LGMF936;C
PC20312
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343165
KC343407
KC343649
KC343891KC344133
–
LGMF915;C
PC20291
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343166
KC343408
KC343650
KC343892KC344134
–
LGMF939;C
PC20315
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
S.A.V.P
ileggi
KC343167
KC343409
KC343651
KC343893KC344135
–
LGMF945;C
PC20321
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343168
KC343410
KC343652
KC343894KC344136
Dia
porth
e pa
divar.p
adi
D. d
ecor
tican
sCBS114200;UPSC2569
Pru
nus
padu
s R
osac
eae
Sweden
K.&
L.H
olm
KC343169
KC343411
KC343653
KC343895KC344137
D
. val
sifo
rmis
CBS114649;UPSC3496
Aln
us g
lutin
osa
Bet
ulac
eae
Sweden
K.&
L.H
olm
KC343170
KC343412
KC343654
KC343896KC344138
Dia
porth
e pa
rane
nsis,sp.nov.
–CBS133184;LGMF929;
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343171
KC343413
KC343655
KC343897KC344139
CPC20305(ex-type)
Dia
porth
e pe
rjunc
ta
D. p
erju
ncta
CBS109745;ARSEF3461;
Ulm
us g
labr
a U
lmac
eae
Austria
A.Y.R
ossm
an
KC343172
KC343414
KC343656
KC343898KC344140
AR3461(ex-epitype)
Dia
porth
e pe
rsea
e,com
b.nov.
P. p
erse
ae
CBS151.73
Per
sea
grat
issi
ma,
you
ng fr
uit
Laur
acea
eNetherlands
E.Laville
KC343173
KC343415
KC343657
KC343899KC344141
A
ntill
es
Dia
porth
e ph
aseo
loru
m
P. o
lear
iae
CBS113425
Ole
ariacf.
rani
A
ster
acea
eNew
Zealand
G.J.M.Verkley
KC343174
KC343416
KC343658
KC343900KC344142
D
. pha
seol
orum
CBS116019;STA
M30
Cap
eron
ia p
alus
tris
Eup
horb
iace
ae
USA:M
ississippiA.M
engistu
KC343175
KC343417
KC343659
KC343901KC344143
D
. pha
seol
orum
CBS116020;STA
M31
Ast
er e
xilis
A
ster
acea
eUSA:M
ississippiA.M
engistu
KC343176
KC343418
KC343660
KC343902KC344144
D
iapo
rthesp.
CBS127465;G
JS83-379
Act
inid
ia c
hine
nsis
, rot
ting
fruit
Act
inid
iace
ae
New
Zealand
S.R.P
ennycook
KC343177
KC343419
KC343661
KC343903KC344145
D
. mel
onis
CBS257.80
––
–L.Beraha
KC343178
KC343420
KC343662
KC343904KC344146
D
iapo
rthesp.
LGMF927;C
PC20303
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343179
KC343421
KC343663
KC343905KC344147
D
iapo
rthesp.
LGMF941;C
PC20317
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343180
KC343422
KC343664
KC343906KC344148
Dia
porth
e ps
eudo
man
gife
rae,
P.
man
gife
rae
CBS101339(ex-type)
Man
gife
ra in
dica
A
naca
rdia
ceae
Dom
inican
P.deLeeuw
KC343181
KC343423
KC343665
KC343907KC344149
sp.nov.
Republic
P.
man
gife
rae
CBS388.89
Man
gife
ra in
dica
, pee
l of f
ruit
Ana
card
iace
ae
Mexico
–KC343182
KC343424
KC343666
KC343908KC344150
Dia
porth
e ps
eudo
phoe
nici
cola
, P.
man
gife
rae
CBS176.77
Man
gife
ra in
dica
, sho
win
g di
ebac
k A
naca
rdia
ceae
Iraq
M.S.A.Al-M
omen
KC343183
KC343425
KC343667
KC343909KC344151
sp.nov.
P.
pho
enic
icol
aCBS462.69(ex-type)
Pho
enix
dac
tylif
era,
dea
d to
ps o
f A
reca
ceae
Spain
H.A.vanderAa
KC343184
KC343426
KC343668
KC343910KC344152
gr
een
leav
es
Dia
porth
e pu
stul
ata
D. p
ustu
lata
CBS109742;AR3430
Ace
r pse
udop
lata
nus
Ace
race
ae
Austria
A.Y.R
ossm
an
KC343185
KC343427
KC343669
KC343911
KC344153
D
. pus
tula
ta
CBS109760;AR3535
Ace
r pse
udop
lata
nus
Ace
race
ae
Austria
A.Y.R
ossm
an
KC343186
KC343428
KC343670
KC343912KC344154
D
. pad
iCBS109784;AR3419
Pru
nus
padu
s R
osac
eae
Austria
A.Y.R
ossm
an
KC343187
KC343429
KC343671
KC343913KC344155
Dia
porth
e ra
onik
ayap
orum
,sp.nov.–
CBS133182;LGMF923;
Spo
ndia
s m
ombi
n, e
ndop
hytic
in le
af
Ana
card
iace
ae
Brazil
K.R
odriguez
KC343188
KC343430
KC343672
KC343914KC344156
CPC20299(ex-type)
Dia
porth
e rh
oina
D
. rho
ina
CBS146.27
Rhu
s to
xico
dend
ron
Ana
card
iace
ae
–L.E.W
ehmeyer
KC343189
KC343431
KC343673
KC343915KC344157
Dia
porth
e sa
ccar
ata,com
b.nov.
P. s
acca
rata
CBS116311;STE
-U3743;
Pro
tea
repe
ns, c
anke
rs
Pro
teac
eae
SouthAfrica
S.D
enman
KC343190
KC343432
KC343674
KC343916KC344158
CPC3743(ex-type)
Dia
porth
e sc
hini,sp.nov.
–CBS133181;LGMF921;
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343191
KC343433
KC343675
KC343917KC344159
CPC20297(ex-type)
–
LGMF910;C
PC20286
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343192
KC343434
KC343676
KC343918KC344160
Dia
porth
e sc
lero
tioid
es
P. s
cler
otio
ides
CBS296.67;ATC
C18585;
Cuc
umis
sat
ivus
, roo
t C
ucur
bita
ceae
Netherlands
H.A.vanderKesteren
KC343193
KC343435
KC343677
KC343919KC344161
IMI151828(ex-type)
P.
scl
erot
ioid
es
CBS710.76;PD76/674
Cuc
umis
sat
ivus
, roo
t C
ucur
bita
ceae
Netherlands
–KC343194
KC343436
KC343678
KC343920KC344162
Dia
porth
e sc
obin
a P.
sco
bina
CBS251.38
Frax
inus
exc
elsi
or, l
ivin
g an
d de
ad tw
ig
Ole
acea
eUK:S
cotland
J.A.M
acDonald
KC343195
KC343437
KC343679
KC343921KC344163
Dia
porth
e so
jae
P. lo
ngic
olla
CBS100.87
Gly
cine
soj
a Fa
bace
ae
Italy
P.Giunchi
KC343196
KC343438
KC343680
KC343922KC344164
P.
long
icol
la
CBS116017;DP0508;S
TAM28
Eup
horb
ia n
utan
s E
upho
rbia
ceae
USA:M
ississippiA.M
engistu
KC343197
KC343439
KC343681
KC343923KC344165
P.
long
icol
la
CBS116023;STA
M35
Gly
cine
max
Fa
bace
ae
USA:M
ississippiA.M
engistu
KC343198
KC343440
KC343682
KC343924KC344166
P.
long
icol
la
CBS127267;PL4
Gly
cine
max
, ste
m
Faba
ceae
Croatia
K.V
randečić
KC343199
KC343441
KC343683
KC343925KC344167
D
. pha
seol
orum
var.
CBS180.55;ATC
C12050;C
ECT2024;
Gly
cine
soj
a, m
atur
e st
em
Faba
ceae
–A.A.H
ildebrand
KC343200
KC343442
KC343684
KC343926KC344168
so
jae
Alfaro245
D
. pha
seol
orum
var.
CBS659.78;NRRL13656
Gly
cine
soj
a, s
eedl
ing
Faba
ceae
USA
J.Marcinkow
ska
KC343201
KC343443
KC343685
KC343927KC344169
so
jae
6 Persoonia–Volume31,2013
Dia
porth
esp.1
D. p
hase
olor
um
CBS119639;B11861
Man,abscess
–Germany
K.P
lechulla
KC343202
KC343444
KC343686
KC343928KC344170
D
iapo
rthesp.
LGMF947;C
PC20323
Gly
cine
max
, see
d Fa
bace
ae
Brazil
A.Almeida
KC343203
KC343445
KC343687
KC343929KC344171
Dia
porth
esp.2
Dia
porth
esp.
LGMF932;C
PC20308
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343204
KC343446
KC343688
KC343930KC344172
Dia
porth
esp.3
P. c
onor
um
CBS287.29
Pse
udot
suga
men
zies
ii P
inac
eae
UK:S
cotland
G.G.H
ahn
KC343205
KC343447
KC343689
KC343931KC344173
Dia
porth
esp.4
Dia
porth
esp.
LGMF944;C
PC20320
May
tenu
s ili
cifo
lia, e
ndop
hytic
in p
etio
le
Cel
astra
ceae
Brazil
R.R.G
omes
KC343206
KC343448
KC343690
KC343932KC344174
Dia
porth
esp.5
–CBS125575
Ace
r opa
lus
Ace
race
ae
Italy
W.Jaklitsch
KC343207
KC343449
KC343691
KC343933KC344175
Dia
porth
esp.6
P. p
ittos
pori
CBS115584;HKUCC7784;AT7
Mae
sa p
erla
rius,
frui
t M
yrsi
nace
ae
HongKong
K.D.H
yde
KC343208
KC343450
KC343692
KC343934KC344176
P.
pitt
ospo
riCBS115595;HKUCC10129
Mae
sa p
erla
rius,
frui
t M
yrsi
nace
ae
HongKong
K.D.H
yde
KC343209
KC343451
KC343693
KC343935KC344177
Dia
porth
esp.7
P. a
naca
rdii
CBS458.78
Ana
card
ium
occ
iden
tale
A
naca
rdia
ceae
India
H.C.G
ovindu
KC343210
KC343452
KC343694
KC343936KC344178
Dia
porth
esp.8
Dia
porth
esp.
LGMF925;C
PC20301
Asp
idos
perm
a to
men
tosu
m
Apo
cyna
ceae
Brazil
K.R
odriguez
KC343211
KC343453
KC343695
KC343937KC344179
Dia
porth
e st
ictic
a,com
b.nov.
P. s
tictic
aCBS370.54
Bux
us s
empe
rvire
ns, d
ead
twig
B
uxac
eae
Italy
M.R
ibaldi
KC343212
KC343454
KC343696
KC343938KC344180
Dia
porth
e su
bord
inar
ia,com
b.nov.
P. s
ubor
dina
ria
CBS101711
Pla
ntag
o la
nceo
lata
, bla
cken
ed s
eed
Pla
ntag
inac
eae
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Zealand
B.Alexander
KC343213
KC343455
KC343697
KC343939KC344181
P.
sub
ordi
naria
CBS464.90
Pla
ntag
o la
nceo
lata
, sta
lk
Pla
ntag
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R.S
hivas
KC343214
KC343456
KC343698
KC343940KC344182
Dia
porth
e te
com
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P. te
com
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CBS100547
Tabe
buiasp.,m
ycocecidiumcausedby
Big
noni
acea
eBrazil
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KC343215
KC343457
KC343699
KC343941KC344183
P
roso
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um te
com
icol
aD
iapo
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tere
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lii,sp.nov.
–CBS133180;LGMF914;
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343216
KC343458
KC343700
KC343942KC344184
CPC20290(ex-type)
–
LGMF907;C
PC20283
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343217
KC343459
KC343701
KC343943KC344185
–
LGMF909;C
PC20285
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343218
KC343460
KC343702
KC343944KC344186
–
LGMF913;C
PC20289
Sch
inus
tere
bint
hifo
lius,
end
ophy
tic in
leaf
A
naca
rdia
ceae
Brazil
J.Lima
KC343219
KC343461
KC343703
KC343945KC344187
Dia
porth
e to
xica
D
. tox
ica
CBS534.93;ATC
C96741(ex-type)
Lupi
nus
angu
stifo
lius,
ste
m
Faba
ceae
WesternAustraliaJ.B.N
unn
KC343220
KC343462
KC343704
KC343946KC344188
D
. tox
ica
CBS535.93
Lupi
nussp.
Faba
ceae
WesternAustraliaP.M.W
illiamson
KC343221
KC343463
KC343705
KC343947KC344189
D
. tox
ica
CBS546.93
Lupi
nussp.,stem
Fa
bace
ae
WesternAustraliaP.M.W
illiamson
KC343222
KC343464
KC343706
KC343948KC344190
Dia
porth
e va
ccin
ii D
. vac
cini
iCBS118571;G.C.A.Dvacc
Vacc
iniu
m c
orym
bosu
m
Eric
acea
eUSA:M
ichigan
G.C.Adams
KC343223
KC343465
KC343707
KC343949KC344191
P.
vac
cini
iCBS122112;FAU474
Vacc
iniu
m m
acro
carp
on
Eric
acea
eUSA:N
ewJerseyL.Carris
KC343224
KC343466
KC343708
KC343950KC344192
P.
vac
cini
iCBS122114;FAU634
Vacc
iniu
m c
orym
bosu
m
Eric
acea
eUSA:M
ichigan
D.C.R
amsdell
KC343225
KC343467
KC343709
KC343951KC344193
P.
vac
cini
iCBS122115;FAU590
Vacc
iniu
m c
orym
bosu
m
Eric
acea
eUSA:M
ichigan
D.C.R
amsdell
KC343226
KC343468
KC343710
KC343952KC344194
P.
vac
cini
iCBS122116;DF5022
Vacc
iniu
m c
orym
bosu
m
Eric
acea
eUSA:N
orth
D.F.Farr
KC343227
KC343469
KC343711
KC343953KC344195
C
arol
ina
D
. vac
cini
iCBS160.32;IFO32646(ex-type)
Oxy
cocc
us m
acro
carp
os
Eric
acea
eUSA:M
assa-
C.L.S
hear
KC343228
KC343470
KC343712
KC343954KC344196
ch
uset
tsD
iapo
rthe
vexa
ns
P. v
exan
sCBS127.14
Sol
anum
mel
onge
na
Sol
anac
eae
USA
L.L.Harter
KC343229
KC343471
KC343713
KC343955KC344197
Dia
porth
e vi
ticol
a P.
con
trove
rsa
CBS100170
Frax
inus
exc
elsi
or, l
eaf s
pot
Ole
acea
eNetherlands
H.A.vanderAa
KC343230
KC343472
KC343714
KC343956KC344198
D
. auc
ubae
CBS106.95
Auc
uba
japo
nica
, bra
nche
s an
d tw
igs
Auc
ubac
eae
Netherlands
G.J.M.Verkley
KC343231
KC343473
KC343715
KC343957KC344199
D
. med
usae
aCBS109492
Labu
rnum
ana
gyro
ides
Fa
bace
ae
Austria
A.Y.R
ossm
an
KC343232
KC343474
KC343716
KC343958KC344200
D
. par
dalo
ta
CBS109768;AR3478
Epi
lobi
um a
ngus
tifol
ium
O
nagr
acea
eCanada
M.B
arr
KC343233
KC343475
KC343717
KC343959KC344201
D
. viti
cola
CBS113201;STE
-U5683;
Vitis
vin
ifera
Vitaceae
Portugal
A.J.L.P
hillips
KC343234
KC343476
KC343718
KC343960KC344202
CPC5683(ex-epitype)
D
. viti
cola
CBS114011;CPC2677
Vitis
vin
ifera
Vi
tace
ae
Portugal
A.J.L.P
hillips
KC343235
KC343477
KC343719
KC343961KC344203
D
. circ
umsc
ripta
CBS114436;UPSC2960
Sam
bucu
scf.r
acem
osa
Cap
rifol
iace
ae
Sweden
K.&
L.H
olm
KC343236
KC343478
KC343720
KC343962KC344204
D
. med
usae
aCBS266.85;PD85/25
Ros
a ru
gosa
R
osac
eae
Netherlands
G.H.B
oerema
KC343237
KC343479
KC343721
KC343963KC344205
D
. woo
dii
CBS312.91
Lupi
nus
arbo
reus
, dea
d st
em
Faba
ceae
Netherlands
H.A.vanderAa&
KC343238
KC343480
KC343722
KC343964KC344206
F.Meurs
P.
sal
icin
aCBS446.62
Sal
ixsp.,twig
Sal
icac
eae
Netherlands
G.H.B
oerema
KC343239
KC343481
KC343723
KC343965KC344207
D
. woo
dii
CBS449.82
Lupi
nussp.,deadstem
Faba
ceae
Netherlands
H.A.vanderAa
KC343240
KC343482
KC343724
KC343966KC344208
P.
dip
saci
CBS502.85
Dip
sacu
s fu
llonu
m, d
ead
stem
D
ipsa
cace
ae
Netherlands
H.A.vanderAa
KC343241
KC343483
KC343725
KC343967KC344209
P.
asp
hode
lina
CBS759.95
Asp
hode
lus
albu
s, 1
-yr-
old
stem
s A
spho
dela
ceae
France
G.J.M.Verkley
KC343242
KC343484
KC343726
KC343968KC344210
D
. auc
ubae
CBS794.96
Auc
uba
japo
nica
A
ucub
acea
eUK
G.J.M.Verkley
KC343243
KC343485
KC343727
KC343969KC344211
Dia
porth
e w
oodi
i D
. woo
dii
CBS558.93
Lupi
nussp.,stem
Fa
bace
ae
WesternAustraliaP.M.W
illiamson
KC343244
KC343486
KC343728
KC343970KC344212
Dia
porth
e w
oolw
orth
ii D
. woo
lwor
thii
CBS148.27
Ulm
us a
mer
ican
a U
lmac
eae
–L.E.W
ehmeyer
KC343245
KC343487
KC343729
KC343971KC344213
Dia
porth
ella
cor
ylin
a–
CBS121124;AR4131
Cor
ylussp.,dyingstems
Cor
ylac
eae
China:Fuyuan
L.N.Vassiljeva
KC343004
KC343246
KC343488
KC343730KC343972
1 AR:C
ollectionofA.Y.R
ossm
an;ATC
C:A
mericantypeculturecollection;CBS:C
BSFungalB
iodiversityCentre,U
trecht,TheNetherlands;CECT:ColeccionEspañoladeCultivosTipo,UniversityofValencia,Valencia,Spain;C
PC:C
ollectionPedroCrous,housedatCBS;D
AOM:
PlantResearchInstitute,D
epartmentofAgriculture(M
ycology),O
ttawa,Canada;HKUCC:U
niversityofH
ongKongCultureCollection,DepartmentofE
cologyandBiodiversity,H
ongKong,China;IFO
:InstituteforF
ermentation,Osaka,Japan;IMI:InternationalM
ycologicalInstitute,
CABI-B
ioscience,Egham
,Bakeham
Lane,U.K.;LG
MF:CulturecollectionofLaboratoryofGeneticsofMicroorganism
s,FederalUniversityofP
arana,Curitiba,B
razil;MUCL:UniversitéCatholiquedeLouvain,Louvain-la-Neuve,B
elgium
;NRRL:NationalC
enterforAgricultural
UtilizationResearch,Peoria,Illinois,U
.S.A.;PD:P
lantProtectionService,W
ageningen,TheNetherlands;STE
-U:S
tellenboschUniversityculturecollections,S
outhAfrica;U
PSC:FungalC
ultureCollectionattheBotanicalMuseum,U
ppsalaUniversity,U
ppsala,S
weden
2 TU
B:partialbeta-tubulingene;CAL:partialcalmodulingene;HIS:partialhistoneH3gene;ITS
:internaltranscribedspacerregionsofthenrDNAandintervening5.8S
nrDNA;TEF1:partialtranslationelongationfactor1-alphagene.
Tabl
e 1(cont.)
Spe
cies
O
rigin
al n
ame
Stra
in1
Isolationsource
Hostfam
ily
Locality
Collector
GenBa
nkAccessionnum
bers(ITS
,CAL
,HIS,TEF
1,TUB)
2
7R.R.Gomesetal.:Diaporthe
(1933)totheconclusionthathost-associationwasnot infor-mative enough in Diaporthe, thereby reducing the number of speciesfrom650toonly70inthegenus.However,thisrevisionwas based strictly on morphological characters of the Diaporthe sexual state, and connections to the Phomopsis asexual states (priortomolecularanalyses)hadbeenidentifiedonlyin20%ofthespecies(Wehmeyer1933).Although the classificationofDiaporthe has been on-going, speciesarepresentlybeingredefinedbasedonacombina-tion of morphological, cultural, phytopathological, mating type andDNAsequencedata(Rehner&Uecker1994,Zhangetal.1998,Mostertetal.2001a, Farretal.2002,Santosetal.2010).However,evenwhenusingacombinationofmorphologicalandmolecular data, the delimitation of species within the genus Diaportheonlyprovedsatisfactoryoncemulti-geneDNAse-quencedataweregenerated (Castlebury&Mengistu2006,vanRensburgetal.2006,Santosetal.2010,Udayangaetal.2012),sincethisaddsvaluableinformationintheresolutionofcomplexevolutionary relationships.Theaimsof thepresentstudywerethusto:1)provideamulti-genephylogenyforthegenus Diaporthebasedonalargesetofwell-identifiedculturesdepositedintheCBSculturecollection;2)toidentifypotentialisolatesforepitypification,therebyfixingtheapplicationofpre- viouslyestablishednames;3)tolinkDiaporthe names to their Phomopsis asexual states;and4) to identifyacollectionofmostly sterile endophytic Diaporthe strains isolated from several medicinalhostsinBrazil.
MATERIALS AND METHODS
IsolatesInthepresentstudyweanalysed243Diaportheisolates(Table1),aswellastheoutgroupDiaporthella corylina.Isolateswereobtained from several sources, including 40 endophytic strains isolatedfrommedicinalplantsinBrazil(LabGeM/UFPRcollec-tion,Curitiba,Brazil),andthreeisolatesfromtheEMBRAPA-SOJAcollection,Londrina,Brazil.Afurther199isolateswereobtained from theCBS-KNAWFungal Biodiversity Centre(CBS),ortheworkingcollectionofP.W.Crous(CPC)housedatCBS.
DNA isolation, amplification and phylogenetic analysisColonieswerecultivatedon2%potato-dextroseagar(PDA),andgenomicDNAextractionwasundertakenusingtheUltra-Clean™MicrobialDNAKit(MOBio,Carlsbad,CA,USA)ac-cordingtomanufacturer’sinstructions. Using20isolates,wescreenednineloci,ofwhichthefivemoreinformativelociwereselectedformulti-geneanalyses.TheprimersITS5andITS4(Whiteetal.1990)wereusedtoam-plifytheinternaltranscribedspacerregion(ITS)ofthenuclear ribosomalRNAgeneoperon,includingthe3’endofthe18SnrRNA, thefirst internal transcribedspacer region, the5.8SnrRNAgene; the second internal transcribed spacer regionandthe5’endofthe28SnrRNAgene.TheprimersEF1-728FandEF1-986R(Carbone&Kohn1999)wereusedtoamplifypartofthetranslationelongationfactor1-αgene(TEF1)andtheprimersACT-512FandACT-783R(Carbone&Kohn1999)wereusedtoamplifypartoftheactingene(ACT).Theprim-ersGpd1-LMandGpd2-LM(Myllysetal.2002)wereusedtoamplify part of the glyceraldehyde-3-phosphate dehydrogenase (GPDH)gene,andpartofthecalmodulin(CAL)genewasse-quencedusingtheprimersCAL-228FandCAL-737R(Carbone&Kohn1999).TheprimersCYLH3F(Crousetal.2004b)andH3-1b(Glass&Donaldson1995)wereusedtoamplifypartofthehistoneH3(HIS)gene,andtheprimersT1(O’Donnell&Cigelnik1997)andBt-2b(Glass&Donaldson1995)toamplify
partoftheβ-tubulingene(TUB).TheprimersNMS1andNMS2(Lietal.1994)wereusedtoamplifyaninternalregionofthemitochondrialSSU(mtSSU).ThepartiallargesubunitnrDNA(LSU)wassequencedusingtheprimersLSU1Fd(Crousetal.2009a)andLR5(Vilgalys&Hester1990).Amplificationreactionshadatotalreactionvolumeof12.5μLwhich was composed of 1×PCRbuffer(BiolineGmbH,Lucken- walde,Germany),5.6%DMSO(v/v),20μMdNTPs,0.2μMofeach forwardand reverseprimers,0.25UofBioTaqTaq DNApolymerase(BiolineGmbH,Luckenwalde,Germany),and10ngofgenomicDNA.PCRconditionswerethesameforallloci,exceptfortheMgCl2
concentration:2mMMgCl2 for the
genesLSUandTEF1,1.5mMMgCl2 forthegenesACT,GPDH,
mtSSU,ITSandTUB,and1mMMgCl2forCALandHISgenes.ThePCRconditionswere:startstepof2minat94°C,followedby40cyclesof30sat94°C,1minatadequateannealingtemperature,and1minat72°C,followedbyafinishingstepof3minat72°Candacooldownstepto4°C.Theannealingtemperaturevariedforeachgene:61°C(ACT,GPDH,mtSSU);58°C(CAL,ITS,HIS);55°C(TEF1,TUB)and48°C(LSU).However, someof theseprimer pairs failed to amplifywithsome isolates included in this study, and therefore additional combinationswereused.Theamplificationreactionandcycleconditions were the same except the annealing temperature and MgCl2
concentration.FortheamplificationofTEF1withprimers
EF1-728FandEF2(O’Donnelletal.1998),52°Cand2mMMgCl2;TUBwithprimersT1(O’Donnell&Cigelnik1997)andCYLTUB1R(Crousetal.2004b),50°Cand1mMMgCl2;CALwithprimersCAL-228FandCAL2Rd(Quaedvlieg etal.2011,Groenewaldetal.2013),58°Cand1mMMgCl2.Ampliconswere sequencedusingbothPCRprimerswith aBigDyeTerminatorCycleSequencingKitv.3.1(AppliedBiosys-tems,FosterCity,CA,USA)accordingtothemanufacturer’sin-structions,andsequenceswereanalysedonanABIPrism3700DNASequencer(Perkin-Elmer,Norwalk,FosterCity,CA,USA).TheconsensussequenceswerevisuallyinspectedusingMEGAv.5software(Tamuraetal.2011).ThealignmentofobtainedsequenceswasperformedusingtheonlineMAFFTinterface(Katoh&Toh2008;http://mafft.cbrc.jp/alignment/server).ForthephylogeneticanalysesbasedonMaximumLikelihoodand Bayesian inference, we chose the best evolutionary models foreachdatapartitionusingthesoftwareMrModelTestv.2.3(Nylander2004).MrBayesv.3.1.1(Ronquist&Huelsenbeck2003)wasusedtogeneratethephylogenetictreesunderopti-malcriteriaperdatapartition. Theheatingparameterwassetat0.3andtheMarkovChainMonteCarlo(MCMC)analysisoffourchains was started in parallel from a random tree topology and lasteduntiltheaveragestandarddeviationofsplitfrequenciescamebelow0.01.Treesweresavedeach10000generationsandtheresultingphylogenetic tree(Fig.1)wasprintedwithGeneiousv.5.5.4(Drummondetal.2011)andthelayoutofthetreewasdoneinAdobeIllustratorv.CS5.1.Diaporthella corylina (CBS121124) was used as outgroup in the phylogenetic analyses based on its position as sister family in Diaporthales (Vasilyevaetal.2007).NewsequencesgeneratedinthisstudyweredepositedinNCBIsGenBanknucleotidedatabase(www.ncbi.nlm.nih.gov;Table1)andthealignmentandphylogenetictreeinTreeBASE(studyS13943;www.treebase.org).
Locus resolution and SNP detectionNeighbour-joininganalysesusingthegeneraltime-reversiblesubstitution model were applied to each data partition individu-ally to check the stability and robustness of each species clade under each dataset usingPAUPv.4.0b10 (Swofford 2003)(TreeBASEstudyS13943).Alignmentgapswere treatedasmissingdataandallcharacterswereunorderedandofequal
8 Persoonia–Volume31,2013
CBS 116023 Phomopsis longicolla Glycine max USA CBS 659.78 D. phaseolorum var. sojae Glycine soja USA CBS 116017 Phomopsis longicolla Euphorbia nutans USA CBS 100.87 Phomopsis longicolla Glycine soja Italy CBS 127267 Phomopsis longicolla Glycine max Croatia CBS 180.55 D. phaseolorum var. sojae Glycine soja Unknown CBS 119639 “Diaporthe phaseolorum” Man, abscess Germany LGMF947 Glycine max Brazil CBS 124654 Phomopsis convolvuli Convolvulus arvensis Turkey LGMF941 Maytenus ilicifolia Brazil LGMF927 Maytenus ilicifolia Brazil CBS 257.80 “Diaporthe melonis” Unknown Unknown CBS 116019 Diaporthe phaseolorum Caperonia palustris USA CBS 127465 Diaporthe sp. Actinidia chinensis New Zealand CBS 113425 Phomopsis oleariae Olearia cf. rani New Zealand CBS 116020 Diaporthe phaseolorum Aster exilis USA LGMF935 Maytenus ilicifolia Brazil LGMF928 Maytenus ilicifolia Brazil LGMF919 Schinus terebinthifolius Brazil LGMF934 Maytenus ilicifolia Brazil LGMF937 Maytenus ilicifolia Brazil LGMF911 Schinus terebinthifolius Brazil CBS 133811 = LGMF916 Schinus terebinthifolius Brazil LGMF948 Glycine max Brazil CBS 199.39 Diaporthe conorum Unknown Italy CBS 230.52 Diaporthe citri Citrus sinensis Suriname LGMF946 Glycine max Brazil CBS 507.78 Diaporthe melonis Cucumis melo USA CBS 435.87 D. phaseolorum var. sojae Glycine soja Indonesia CBS 122.21 Diaporthe batatas Ipomoea batatas USA
LGMF909 Schinus terebinthifolius Brazil LGMF907 Schinus terebinthifolius Brazil CBS 133180 = LGMF914 Schinus terebinthifolius Brazil LGMF913 Schinus terebinthifolius Brazil CBS 100547 “Phomopsis tecomae” Tabebuia sp. Brazil
LGMF910 Schinus terebinthifolius Brazil CBS 133181 = LGMF921 Schinus terebinthifolius Brazil
CBS 127.14 Phomopsis vexans Solanum melongena Unknown CBS 481.92 Phomopsis hordei Hordeum vulgare Norway
CBS 592.81 Diaporthe helianthi Helianthus annuus Serbia CBS 344.94 Diaporthe helianthi Helianthus annuus Unknown
CBS 143.27 Diaporthe megalospora Sambucus canadensis Unknown
100
100
100
100
10058
100
100
99
94
100
99
100
100
100
63
100
100
100
10071
97
100
10010081
100
100100
98
0.2
D. phaseolorum 4
2Diaporthe sp. 1
1Diaporthe sojae
Diaporthe terebinthifolii 9
Diaporthe endophytica 5
3D. convolvuli
Diaporthe citri 6
Diaporthe vexans 1213 Diaporthe hordei
Diaporthe schini
Diaporthe tecomae
11
10
8Diaporthe batatas
Diaporthe melonis 7
15D. megalospora
14 Diaporthe helianthi
Fig. 1Consensusphylogramof22104treesresultingfromaBayesiananalysisofthecombined5-genesequencealignment.Cladesarenumberedontheright of the boxes and Diaporthespeciesnamesinpurplereflectnewcombinationsandinrednewspecies.Strainaccessionnumbersarefollowedbytheoriginalspeciesname(black,whenapplicable),theisolationsource(green)andcountryoforigin(blue).Accessionnumbersandnamesinbold represent strainsknowntobeex-typestrainsorareconsideredtobeauthenticforthespecies.Reddotsindicatestrainsfrommedicinalplantsandyellowdotsfromhumans.Bayesianposteriorprobabilitiesareshownatthenodesandthescalebarrepresentstheexpectedchangespersite.ThetreewasrootedtoDiaporthella corylina(strainCBS121124).
weight.Any tieswere broken randomlywhenencountered.Therobustnessofthetreesobtainedwasevaluatedby1000bootstrapreplications(Hillis&Bull1993).Inthepresentstudy,boththeanalysisofthecombinedalignment(Fig.1)andoftheindividuallociwereusedtodeterminethespeciesboundaries.For each clade in the combined analysis, the position of the members of that clade was determined in the phylogenetic tree obtained from each of the individual loci to check whether these members still represent a single clade in the individual genetree.Inthiswaytherobustnessofagivencladecould
be evaluated together with the posterior probability value of thatclade.Aspecieswasonlycountedifitwasdistinctfromits closest relatives and the species clade contained all the associatedstrains.Unique fixed nucleotide positions are used to characteriseanddescribeseveralsterilespecies(seeapplicablespeciesnotes).Foreachsterilespeciesthatwasdescribed,theclos-estphylogeneticneighbour(s)wereselectedfromFig.1andthisfocuseddatasetwassubjectedtoSNPanalyses.Thesesinglenucleotidepolymorphisms(SNPs)weredeterminedfor
9R.R.Gomesetal.:Diaporthe
CBS 109490 “Diaporthe arctii” Ambrosia trifida USA CBS 501.90 Diaporthe angelicae Heracleum sphondylium France CBS 111591 Diaporthe angelicae Heracleum sphondylium Austria CBS 111592 Diaporthe angelicae Heracleum sphondylium Austria CBS 123215 Diaporthe angelicae Foeniculum vulgare Portugal CBS 344.86 Phomopsis asteriscus Eryngium maritimum France CBS 100871 Phomopsis foeniculi Foeniculum vulgare Italy CBS 101711 Phomopsis subordinaria Plantago lanceolata New Zealand CBS 464.90 Phomopsis subordinaria Plantago lanceolata South Africa CBS 136.25 Diaporthe arctii Arctium sp. Unknown CBS 117499 Phomopsis cuppatea Aspalathus linearis South Africa CBS 123213 Diaporthe lusitanicae Foeniculum vulgare Portugal CBS 123212 Diaporthe lusitanicae Foeniculum vulgare Portugal LGMF943 Maytenus ilicifolia Brazil
CBS 127271 Diaporthe novem Glycine max Croatia CBS 354.71 Diaporthe pardalota Polygonatum odoratum Romania CBS 127269 Diaporthe novem Glycine max Croatia CBS 127270 Diaporthe novem Glycine max Croatia
CBS 133812 = LGMF906 Schinus terebinthifolius Brazil LGMF912 Schinus terebinthifolius Brazil LGMF920 Schinus terebinthifolius Brazil LGMF933 Maytenus ilicifolia Brazil LGMF918 Schinus terebinthifolius Brazil LGMF940 Maytenus ilicifolia Brazil
LGMF908 Schinus terebinthifolius Brazil LGMF917 Schinus terebinthifolius Brazil CBS 180.91 Phomopsis ganjae Cannabis sativa USA CBS 505.76 Phomopsis manihot Manihot utilissima Rwanda
CBS 114015 Diaporthe ambigua Pyrus communis South Africa CBS 117167 Diaporthe ambigua Aspalathus linearis South Africa CBS 123211 Diaporthe ambigua Foeniculum vulgare Portugal CBS 123210 Diaporthe ambigua Foeniculum vulgare Portugal CBS 127746 Diaporthe scabra Platanus acerifolia Italy CBS 187.87 “Diaporthe helianthi” Helianthus annuus Italy CBS 194.36 Diaporthe strumella var. longispora Ribes sp. Canada CBS 296.67 Phomopsis sclerotioides Cucumis sativus Netherlands CBS 710.76 Phomopsis sclerotioides Cucumis sativus Netherlands
LGMF932 Maytenus ilicifolia Brazil CBS 133185 = LGMF938 Maytenus ilicifolia Brazil CBS 133182 = LGMF923 Spondias mombin Brazil
CBS 287.29 Phomopsis conorum Pseudotsuga menziesii UK LGMF944 Maytenus ilicifolia Brazil
LGMF915 Schinus terebinthifolius Brazil CBS 133186 = LGMF942 Maytenus ilicifolia Brazil LGMF939 Maytenus ilicifolia Brazil LGMF945 Maytenus ilicifolia Brazil LGMF936 Maytenus ilicifolia Brazil
CBS 133184 = LGMF929 Maytenus ilicifolia Brazil CBS 133183 = LGMF924 Aspidosperma tomentosum Brazil LGMF926 Aspidosperma tomentosum Brazil CBS 125575 Acer opalus Italy
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Diaporthe paranensis 35
Diaporthe oxe 34
Diaporthe sp. 5 37
Diaporthe brasiliensis 36
33 Diaporthe sp. 432 Diaporthe sp. 3
Diaporthe neoarctii 16
19
18
Diaporthe arctii
D. subordinaria
26Diaporthe ambigua
31 Diaporthe raonikayaporum Diaporthe mayteni 30
29 Diaporthe sp. 2
22Diaporthe novem
17D. angelicae
Diaporthe infecunda 23
24 Diaporthe ganjae25 Diaporthe manihotia
Diaporthe cuppatea 20
21 Diaporthe lusitanicae
28 Diaporthe sclerotioides
27 Diaporthe longispora
Fig. 1(cont.)
eachaligneddatapartitionusingDnaSPv.5.00.07(Librado&Rozas2009).
TaxonomyAll descriptions provided are based on colonies sporulating in culture,whichforthemostpartonlyformedtheasexualmorph.Coloniesweresubculturedonto2%tapwateragarsupple-mentedwithsterilepineneedles(PNA;Smithetal.1996),or
autoclaved leaf pieces of Ilex aquifolium, Maytenus ilicifolia or Schinus terebinthifolius,PDA,oatmealagar(OA),and2%maltextractagar(MEA)(accordingtoCrousetal. 2009b),andincubatedat20°Cundera12hnear-ultravioletlight(400–315nm)(SylvaniaBlacklight-Blue,OsramNederlandB.V.,AlphenaandenRijn,TheNetherlands),12hdarkcycle topromotesporulation.Structuresweremountedinclearlacticacid,and50measurements determined for conidia, and 30for other
10 Persoonia–Volume31,2013
CBS 251.38 Phomopsis scobina Fraxinus excelsior UK CBS 137.27 Diaporthe acerina Acer saccharum Unknown CBS 109745 Diaporthe perjuncta Ulmus glabra Austria
CBS 114435 Diaporthe crataegi Crataegus oxyacantha Sweden CBS 126679 Phomopsis amygdali 3B Prunus dulcis Portugal CBS 115620 Phomopsis amygdali Prunus persica USA CBS 120840 Phomopsis amygdali Prunus salicina South Africa CBS 126680 Phomopsis amygdali 55A Prunus dulcis Portugal CBS 111811 Phomopsis amygdali Vitis vinifera South Africa
CBS 129521 Diaporthe acaciigena Acacia retinodes Australia CBS 109742 Diaporthe pustulata Acer pseudoplatanus Austria CBS 109784 Diaporthe padi Prunus padus Austria CBS 109760 Diaporthe pustulata Acer pseudoplatanus Austria
CBS 534.93 Diaporthe toxica Lupinus angustifolius Western Australia CBS 546.93 Diaporthe toxica Lupinus sp. Western Australia CBS 535.93 Diaporthe toxica Lupinus sp. Western Australia
CBS 852.97 “Phomopsis heveae” Hevea brasiliensis Brazil CBS 138.27 Diaporthe beckhausii Viburnum sp. Unknown CBS 122676 Diaporthe cynaroidis Protea cynaroides South Africa CBS 111886 Diaporthe australafricana Vitis vinifera Australia CBS 113487 Diaporthe australafricana Vitis vinifera South Africa CBS 446.62 Phomopsis salicina Salix sp. Netherlands CBS 449.82 “Diaporthe woodii” Lupinus sp. Netherlands CBS 113201 Diaporthe viticola Vitis vinifera Portugal CBS 100170 Phomopsis controversa Fraxinus excelsior Netherlands CBS 759.95 Phomopsis asphodelina Asphodelus albus France CBS 109492 Diaporthe medusaea Laburnum anagyroides Austria CBS 266.85 Diaporthe medusaea Rosa rugosa Netherlands CBS 114011 Diaporthe viticola Vitis vinifera Portugal CBS 114436 Diaporthe circumscripta Sambucus cf. racemosa Sweden CBS 106.95 Diaporthe aucubae Aucuba japonica Netherlands CBS 109768 Diaporthe pardalota Epilobium angustifolium Canada CBS 794.96 Diaporthe aucubae Aucuba japonica UK CBS 502.85 Phomopsis dipsaci Dipsacus fullonum Netherlands CBS 312.91 “Diaporthe woodii” Lupinus arboreus Netherlands
CBS 114434 Diaporthe impulsa Sorbus aucuparia Sweden CBS 141.27 Diaporthe impulsa Sorbus americana Unknown CBS 113830 Diaporthe fibrosa Rhamnus cathartica Sweden CBS 109751 Diaporthe fibrosa Rhamnus cathartica Austria CBS 267.80 Phomopsis viticola Vitis vinifera Italy CBS 114867 Phomopsis viticola Vitis vinifera Turkey CBS 111888 Phomopsis viticola Vitis vinifera USA
CBS 140.27 Diaporthe detrusa Berberis vulgaris Unknown CBS 109770 Diaporthe detrusa Berberis vulgaris Austria CBS 114652 Diaporthe detrusa Berberis vulgaris Sweden CBS 114437 Diaporthe carpini Carpinus betulus Sweden CBS 114649 Diaporthe valsiformis Alnus glutinosa Sweden CBS 114200 Diaporthe decorticans Prunus padus Sweden CBS 148.27 Diaporthe woolworthii Ulmus americana Unknown
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Diaporthe viticola 50
56Diaporthe padi var. padi
51Diaporthe impulsa
52Diaporthe fibrosa
54Diaporthe detrusa
55Diaporthe carpini
Diaporthe beckhausii 47
53Diaporthe ampelina
46Diaporthe cf. heveae 1
49Diaporthe australafricana
48Diaporthe cynaroidis
42Diaporthe amygdali
44Diaporthe pustulata
38 Diaporthe scobina Diaporthe acerina 39
Diaporthe perjuncta 40 Diaporthe crataegi 41
45Diaporthe toxica
43Diaporthe acaciigena
Diaporthe woolworthii 57
Fig. 1(cont.)
structures.The95%confidencelevelsweredetermined,andtheextremesgiven in parentheses.Colonydiametersweredeterminedat25°CindarknessonPDA,OAandMEA.Colonycolours(surfaceandreverse)weredescribedafter14dusingthecolourchartsofRayner(1970).NomenclaturalnoveltiesanddescriptionsweredepositedinMycoBank(www.MycoBank.org;Crousetal.2004a).
RESULTS
DNA sequencing and phylogenetic analysisThemostsuitablegenesforDiaporthe species delimitation in thisstudywere foundtobeCAL,HIS, ITS,TEF1andTUB.Theamplifiedgenomicregionsofthesegenesweremorein-formative, and the combined analysis provided a more robust species identification, fromwhichphylogenetic relationshipscould be inferred.
11R.R.Gomesetal.:Diaporthe
CBS 157.29 Diaporthe nomurai Morus sp. Japan CBS 288.56 Phomopsis gardeniae Gardenia florida Italy CBS 144.27 Diaporthe neilliae Spiraea sp. Unknown CBS 159.47 Diaporthe alnea Alnus sp. Unknown CBS 146.46 Diaporthe alnea Alnus sp. Unknown CBS 587.79 Phomopsis conorum Pinus pentaphylla Japan CBS 338.89 Diaporthe pulla Hedera helix Yugoslavia CBS 116953 Phomopsis fukushii Pyrus pyrifolia New Zealand CBS 129167 Phomopsis sp. 22 Rhododendron sp. Latvia CBS 200.39 Diaporthe nobilis Laurus nobilis Germany CBS 124030 Diaporthe perniciosa Malus pumila New Zealand CBS 113470 Phomopsis castanea Castanea sativa Korea CBS 116954 Phomopsis fukushii Pyrus pyrifolia New Zealand
CBS 122114 Phomopsis vaccinii Vaccinium corymbosum USA CBS 122115 Phomopsis vaccinii Vaccinium corymbosum USA CBS 122116 Phomopsis vaccinii Vaccinium corymbosum USA CBS 122112 Phomopsis vaccinii Vaccinium macrocarpon USA CBS 118571 Diaporthe vaccinii Vaccinium corymbosum USA CBS 495.72 Diaporthe alleghaniensis Betula alleghaniensis Canada
CBS 121004 Diaporthe juglandina Juglans sp. USA CBS 139.27 Diaporthe celastrina Celastrus scandens Unknown CBS 267.32 Phomopsis stictica Unknown Unknown CBS 375.61 Diaporthe perniciosa Malus sylvestris Unknown CBS 445.62 Phomopsis cruciferae Alliaria officinalis Netherlands CBS 283.85 Phomopsis ranojevicii Allium giganteum Netherlands CBS 365.97 Phomopsis cacti Opuntia sp. Netherlands CBS 129168 Phomopsis sp. 23 Rhododendron sp. Latvia CBS 370.67 Phomopsis crustosa Ilex aquifolium Netherlands CBS 287.74 Diaporthe eres Sorbus aucuparia Netherlands CBS 439.82 Phomopsis cotoneastri Cotoneaster sp. UK CBS 110.85 ”Diaporthe arctii” Arctium sp. Netherlands CBS 122.82 Phomopsis skimmiae Skimmia japonica Netherlands CBS 267.55 Phomopsis rudis Laburnum x watereri 'Vossii' Netherlands CBS 250.38 Phomopsis controversa Fraxinus excelsior UK CBS 528.83 Diaporthe seposita Wisteria sinensis Netherlands CBS 841.84 Phomopsis tritici Hordeum sp. Germany CBS 109767 Diaporthe eres Acer campestre Austria CBS 102.81 Diaporthe medusaea Juglans regia Italy CBS 485.96 Phomopsis durandiana Rumex hydrolapathum Netherlands CBS 297.77 Phomopsis osmanthi Osmanthus aquifolium Netherlands CBS 186.37 Diaporthe conorum Picea abies UK CBS 422.50 Phomopsis phaseoli Phaseolus vulgaris Netherlands CBS 688.97 Phomopsis abutilonis Abutilon sp. Netherlands CBS 694.94 Phomopsis crustosa Ilex aquifolium Netherlands CBS 791.68 Phomopsis magnoliicola Magnolia x soulangeana Netherlands CBS 101742 Diaporthe eres Fraxinus sp. Netherlands
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CBS 160.32 Diaporthe vaccinii Oxycoccus macrocarpos USA
.
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62Diaporthe nobilis complex
63Diaporthe vaccinii
Diaporthe eres 67
Diaporthe nomurai 58 Diaporthe gardeniae 59
61Diaporthe alnea
60Diaporthe neilliae
Diaporthe celastrina65Diaporthe juglandina66
64Diaporthe alleghaniensis
Fig. 1(cont.)
Themanuallyadjusted,combined(ITS,TUB,CAL,TEF1andHIS)alignmentfortheBayesiananalysiscontained243iso-lates(includingtheoutgroupsequences)and2435characterswereusedinthephylogeneticanalysis.Thenumberofuniquesitepatternsperdatapartitionwere210,616,242,281and183,respectivelyandwerebasedon466,874,355,316and424alignmentpositions,respectively.BasedontheresultsofMrModeltest,thefollowingpriorsweresetinMrBayesforthedifferentdatapartitions:allpartitionshaddirichletbasefrequen-
cies andGTR+I+Gmodelswith inversegamma-distributedrateswereimplementedforITSandHIS,andHKY+I+Gwithinversegamma-distributedratesforTUB,CALandTEF1.TheBayesian analysis lasted 14735000 generations and theconsensus trees and posterior probabilities were calculated fromthe22104treesleftafterdiscarding7368trees(thefirst25%ofgenerations)forburn-in(Fig.1).Ninety-fivecladesarerecognisedanddiscussedhere.
12 Persoonia–Volume31,2013
CBS 121124 Diaporthella corylina
CBS 114281 Diaporthe decedens Corylus avellana Sweden CBS 109772 Diaporthe decedens Corylus avellana Austria CBS 720.97 “Phomopsis anacardii” Anacardium occidentale East Africa
CBS 589.78 Diaporthe oncostoma Robinia pseudoacacia France CBS 100454 Diaporthe oncostoma Robinia pseudoacacia Germany CBS 109741 Diaporthe oncostoma Robinia pseudoacacia Russia CBS 809.85 Phomopsis crustosa Ilex aquifolium Germany CBS 116311 Phomopsis saccarata Protea repens South Africa
CBS 145.26 Diaporthe hickoriae Carya glabra Unknown CBS 504.72 Phomopsis elaeagni Elaeagnus sp. Netherlands
CBS 370.54 Phomopsis stictica Buxus sempervirens Italy LGMF931 Maytenus ilicifolia Brazil CBS 133813 = LGMF930 Maytenus ilicifolia Brazil LGMF922 Spondias mombin Brazil
CBS 719.96 Phomopsis cinerascens Ficus carica Bulgaria CBS 753.70 Diaporthe sarothamni Spartium junceum Croatia CBS 454.81 “Phomopsis phoenicicola” Chamaerops humilis Greece CBS 171.78 Phomopsis mali f.sp. amygdali Prunus amygdalus Italy CBS 400.48 Phomopsis casuarinae Unknown India CBS 116957 Phomopsis theicola Pyrus pyrifolia New Zealand CBS 287.56 Phomopsis diospyri Diospyros kaki Italy CBS 357.69 Diaporthe seposita Wisteria sinensis Netherlands CBS 187.27 Phomopsis theicola Camellia sinensis Italy CBS 603.88 Phomopsis bougainvilleae Bougainvillea spectabilis Portugal CBS 123208 Diaporthe neotheicola Foeniculum vulgare Portugal CBS 123209 Diaporthe neotheicola Foeniculum vulgare Portugal CBS 111554 Diaporthe foeniculacea Foeniculum vulgare Portugal CBS 111553 Diaporthe foeniculacea Foeniculum vulgare Spain
CBS 115448 Phomopsis pittospori Dichroa febrifuga Hong Kong CBS 681.84 “Phomopsis heveae” Hevea brasiliensis India CBS 114979 Phomopsis pittospori Arenga engleri Hong Kong CBS 101339 Phomopsis mangiferae Mangifera indica Dominican Republic CBS 388.89 Phomopsis mangiferae Mangifera indica Mexico CBS 444.82 Phomopsis eugeniae Eugenia aromatica West Sumatra CBS 129519 Diaporthe musigena Musa sp. Australia CBS 115595 Phomopsis pittospori Maesa perlarius Hong Kong CBS 115584 Phomopsis pittospori Maesa perlarius Hong Kong CBS 151.73 Phomopsis perseae Persea gratissima Netherlands Antilles CBS 535.75 Diaporthe citri Citrus sp. Suriname CBS 161.64 Phomopsis phoenicicola Areca catechu India
CBS 458.78 “Phomopsis anacardii” Anacardium occidentale India CBS 176.77 Phomopsis mangiferae Mangifera indica Iraq CBS 462.69 “Phomopsis phoenicicola” Phoenix dactylifera Spain LGMF925 Aspidosperma tomentosum Brazil
CBS 178.55 D. phaseolorum var. caulivora Glycine soja Canada CBS 127268 Diaporthe caulivora Glycine max Croatia
CBS 162.33 Diaporthe crotalariae Crotalaria spectabilis USA CBS 117168 Diaporthe aspalathi Aspalathus linearis South Africa CBS 117169 Diaporthe aspalathi Aspalathus linearis South Africa CBS 117500 Diaporthe aspalathi Aspalathus linearis South Africa CBS 558.93 Diaporthe woodii Lupinus sp. Western Australia
CBS 146.27 Diaporthe rhoina Rhus toxicodendron Unknown
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68Diaporthe decedens
75Diaporthe inconspicua
Diaporthe foeniculacea 78
Diaporthe cinerascens
Diaporthe chamaeropis
76
77
69Diaporthe anacardii
Diaporthe oncostoma 70
Diaporthe saccarata 71
Diaporthe elaeagni 73
Diaporthe stictica 74
79Diaporthe hongkongensis 80Diaporthe cf. heveae 2
72 Diaporthe hickoriae
82
81
D. pseudomangiferae
83Diaporthe eugeniae
Diaporthe arengae
84Diaporthe musigena
85Diaporthe sp. 6
90Diaporthe sp. 8
87Diaporthe arecae
Diaporthe caulivora 91
86 Diaporthe perseae
94Diaporthe woodii
Diaporthe sp. 7 88
93Diaporthe aspalathi
92Diaporthe crotalariae
Diaporthe rhoina 95
89Diaporthe pseudophoenicicola
2x
Fig. 1(cont.)
Locus resolution and SNP detectionThemtSSUandLSUregionshadveryfewinformativesitesfor the tested strains and were therefore not selected as good markersatspecieslevel.TheACTandGPDHregionswerealsodiscarded as suitable candidates for the multi-gene analyses because of their long branch lengths which made unambigu-
ousalignmentsimpossible.Thesefourlociwerethereforenotusedforfurtheramplificationandsequencingonthecompletedataset.Theremainingfivelocihadvariedsuccessforspe-ciesidentificationandsomephylogeneticlineagesweremorepronetolessvariabilitythanothers.Fifty-eightofthe95spe-
13R.R.Gomesetal.:Diaporthe
cies could be identifiedbyall five loci.The loci are treatedindividually below:
CAL–The locus could distinguish 74 of the 95 species(78%success). It haddifficulty separating: D. endophytica and D. phaseolorum(clades4,5);D. angelicae, D. arctii and D. subordinaria (clades17–19);D. alleghaniensis, D. alnea, D. celastrina, D. eres, D. juglandina, D. neilliae and D. nobilis (clades60–62,64–67);andD. eugeniae, D. musigena, D. perseae, D. pseudomangiferae, D. pseudophoenicicola, Diaporthe sp.6andDiaporthe sp. 7 (clades82–86,88, 89).A singlestrain each of D. angelicae(clade17),D. novem(clade22)and D. terebinthifolii (clade9) clusteredseparate from theotherstrainsofthespecies.
HIS–Thelocuscoulddistinguish84ofthe95species(88%success). It haddifficulty separating: D. australafricana and D. viticola(clades49,50);D. celastrina, D. eres and D. nobilis (clades66,67,62);D. arecae and D. perseae (clades86,87);and D. pseudophoenicicola and Diaporthesp.8(clades89,90).A single strain each of D. endophytica(clade5)andD. tere binthifolii(clade9)clusteredseparatefromtheotherstrainsofthespecies.ThisistheonlylocusthatcandistinguishD. an gelicae(clade17).
ITS–Thelocuscoulddistinguish75ofthe95species(79%success).Ithaddifficultyseparating: D. angelicae, D. arctii and D. subordinaria(clades17–19);D. cynaroides and D. viticola (clades48, 50);D. alnea, D. neilliae and D. nobilis (clades60–62); D. arengae, D. eugeniae and D. pseudomangiferae (clades81–83);D. arecae and D. perseae(clades86,87);and D. aspalathi and D. woodii(clades93,94).Asinglestraineachof D. arecae(clade87),D. inconspicua (clade75),D. novem (clade22)andD. terebinthifolii(clade9),andtwostrainseachof D. impulsa(clade51)andD. infecunda(clade23),clusteredseparatefromtheotherstrainsofthespecies.Thisistheonlylocus that can distinguish D. celastrina (clade17)andD. eres (clade67).
TEF1–The locuscoulddistinguish72of the95species(76%success).Ithaddifficultyseparating:D. tecomae, D. te rebinthifolii(clades9,10);D. angelicae, D. arctii and D. sub ordinaria (clades 17–19);D. australafricana and D. viticola (clades49,50);D. celastrina and D. juglandina (clades65,66);D. eres and D. nobilis(clades67,62);D. chamaeropis, D. cinerascens and D. foeniculaceae (clades 76–78); and D. arengae, D. arecae, D. eugeniae, D. musigena, D. perseae, D. pseudomangiferae, D. pseudophoenicicola, Diaporthesp.6and Diaporthesp.8(clades81–87,89,90).
TUB–The locus could distinguish 84 of the 95 species(88%success).Ithaddifficultyseparating: D. endophytica and D. phaseolorum(clades4,5);D. alleghaniensis, D. celastrina, D. eres, D. juglandina, D. nobilis and D. vaccinii(clades62–67);and D. aspalathi and D. woodii(clades93,94).Asinglestrainof D. angelicae (clade 17) clustered separate from the otherstrainsofthespecies.Thisistheonlylocusthatcandistinguish D. perseae(clade86).
DescriptionsbasedonDNAcharactersareprovidedforthreespeciesintheTaxonomysection,namelyD. endophytica(clade5),D. inconspicua(clade75)andD. infecunda(clade23). Diaporthe endophytica(clade5)wascomparedtoD. phaseolorum (clade4); D. inconspicua(clade75)toD. anacardii(clade69), D. chamaeropis(clade77),D. cinerascens(clade76),D. elaeag ni(clade73),D. foeniculacea(clade78),D. hickoriae(clade72),D. oncostoma(clade70),D. saccarata(clade71)andD. stictica (clade74);andD. infecunda(clade23)toD. angelicae(clade17),D. arctii(clade19),D. cuppatea(clade20),D. lusitanicae(clade21),D. neoarctii(clade16),D. novem(clade22)andD. sub ordinaria(clade18).
TaxonomyThemultigeneanalysesresultedin95well-supportedcladescorrelating to 243 isolates of Diaporthe(Table1,Fig.1).Fif-teen new species are described, nine of which were isolated frommedicinalplants(Aspidosperma tomentosum, Maytenus ilicifolia, Schinus terebinthifolius, Spondias mombin)inBrazil(clades 5, 9, 11, 23, 30, 31, 34, 35 and 36).Twenty-eightclades contain ex-type strains of presently known species, or strains accepted as authentic for the species name or which could be designated as epitypes in the present study, and were thereforewell-resolved(7,8,12,14,17,20–22,24,26–28,40,42,43,45,48–50,63,64,69,71,72,84and91–93).Thesexual-asexual relationship was resolved for several taxa, and is reportedbelow.Newcombinations inDiaporthe are intro-duced below for several Phomopsis names that represented well-resolvedtaxa.Severalpotentialepitypeswereidentifiedduringthisstudy,whicharediscussedbelow.
Diaporthe acaciigena Crous,Pascoe&Jacq.Edwards,Per-soonia26:123.2011
Specimen examined.AustrAliA, Victoria,OtwayRanges,Anglesea,S38°23'21.7"E144°11'12.7",onleavesofAcacia retinodes,16Oct.2009,P.W. Crous, I.G. Pascoe & J. Edwards (holotypeCBSH-20581, ex-typecultureCPC17622=CBS129521).
Notes — Clade 43 contains the ex-type culture of D. acaciigena isolated from Acacia retinodesinAustralia.Thisspeciesis morphologically similar to D. amygdali (clade42)(Crousetal.2011),andcloselyrelatedto D. pustulata (clade44).
Diaporthe acerina(Peck)Sacc.,Syll.Fung.(Abellini)1:611.1882
Basionym.Valsa acerinaPeck,Ann.Rep.N.Y.StateMus.Nat.Hist.28:73.1876.1874.
Specimen examined.unknown, from Acer saccharum,Sept.1927,L.E. Wehmeyer(CBS137.27).
Notes—Clade39 is representedbyD. acerina, isolated from Acer saccharum. This species is genetically similar to D. perjuncta (clade40).ItisknowntooccurinEuropeandNorthAmerica on dead limbs and trunks of Acer pseudoplatanus, A. saccharinum, A. saccharum, A. spicatum, and Acer sp. (Aceraceae)(Spielman 1985,Farretal.1989).
Diaporthe alleghaniensis R.H.Arnold,Canad. J.Bot. 45:787.1967
Specimen examined.CAnAdA, Ontario, on branches of Betula alleghaniensis,June1972,R.H. Arnold(ex-typecultureCBS495.72=ATCC24097=DAOM45776).
Notes—Clade64 contains theex-type strain ofD. alleghaniensis, isolated from Betula alleghaniensis inCanada.Diaporthe alleghaniensis causes canker and dieback of B. alleghaniensis, B. lenta, B. papyrifera and B. pendula in Canada (Arnold1975),buthasalsobeenreportedfromJapan(Farr&Rossman2012).
Diaporthe alnea Fuckel,Jahrb.NassauischenVereinsNaturk.23–24:207.1870(1869–1870)
= Phomopsis alnea Höhn.,Sber.Akad.Wiss.Wien,Math.-naturw.Kl.,Abt.1115:681(33ofrepr.).1906.
Specimens examined.unknown, on Alnussp.,June1946,S. Truter(CBS146.46);onAlnussp.,Aug.1947,S. Truter(CBS159.47).
Notes—Clade61consistsoftwoisolatesfromAlnus(pre-sumablycollectedintheNetherlands).Diaporthe alnea causes dieback of Alnus glutinosa(alder)andA. incana(greyalder).
14 Persoonia–Volume31,2013
IthasbeenreportedfromEurope,RussiaandtheUSA(Munk1957,Oak&Dorset1983,Moricca2002,Mel’niketal.2008,Farr&Rossman2012).
Diaporthe ambigua Nitschke,PyrenomycetesGermanici2:311.1870
Specimens examined. itAly, Sicily, Catania, on Platanus acerifolia, G. Granata(CBS127746=IMI395956);Perugia,onHelianthus annuus,Mar.1987,A. Zazzerini(CBS187.87).–PortugAl,ValeAndeiro,onFoeniculum vulgare, J.M. Santos(CBS123210=Di-C003/10,CBS123211=Di-C002/9).– south AfriCA,WesternCapeProvince,fromPyrus communis, deposited 2002, S. Denman(ex-epitypecultureCBS114015=CPC2657);WesternCape Province, on crown of Aspalathus linearis,15May1997,J.C. Janse van Rensburg(CBS117167=CPC5414).
Notes—Clade26representsD. ambigua, which contains twoisolatespreviouslymisidentifiedasD. scabra (CBS127746)and D. helianthi (CBS187.87), and four isolates of D. ambigua, includingtheex-epitypeculture.Diaporthe ambigua is an important pathogen of Malus domestica, Prunus salicina and Pyrus communis inSouthAfricanfruitorchards.InfectionbyD. ambigua is associated with sunken lesions with longitudinal cracksonaffectedfruittrees.Thefungusrapidlykillsnurseryrootstocks, but also kills mature rootstocks over a longer period oftime(Smitetal.1996).Thisspeciesisalsofoundassaprobeonwildfennel(Santos&Phillips2009).Ithasbeenreportedon Aspalathus linearis (vanRensburgetal.2006),Foeniculum vulgare, Malus domestica (Smitetal.1996,Santos&Phillips2009),Malus sylvestris (Crousetal.2000),Prunus spp. (Smitetal.1996,Mostertetal.2001a),Pyrus communis (Nitschke1867),Pyrus ussuriensis (Tai 1979) andVitis vinifera (vanNiekerketal.2005).Itiswidelydistributed,andisknownfromChina,Cuba(Tai1979),Germany(Nitschke1867),SouthAfrica(Smitetal.1996),UK(Dennis1986)andtheUSA(Washington)(Shaw1973).
Diaporthe ampelina(Berk.&M.A.Curtis)R.R.Gomes, C.Glienke&Crous,comb. nov.—MycoBankMB802922
Basionym.Phoma ampelinaBerk.&M.A.Curtis,Grevillea2,18:81.1873. ≡Phomopsis ampelina(Berk.&M.A.Curt.)Grove,Bull.Misc.Inform.Kew4:184.1919. = Phoma viticolaSacc.,Michelia2:92.1880. ≡Phomopsis viticola(Sacc.)Sacc.,Ann.Mycol.13:118.1915. =Fusicoccum viticolumReddick,CornellUniv.Agric.Exp.Sta.Bull.263:331.1909. ≡ Phomopsis viticola (Reddick) Goid., AttiRealeAccad.Naz.Lincei26:107.1937. =Phomopsis viticolaSacc.var.ampelopsidisGrove,Bull.Misc.Inform.Kew4:183.1919. =Diaporthe neoviticolaUdayanga,Crous&K.D.Hyde,Fung.Diversity56:166.2012(anom.nov.basedonPhoma viticolaSacc.).
Conidiomata pycnidial, eustromatic, subepidermal, brown to black, scattered or aggregated, globose, flask-like to conical, outer surface smooth, convoluted to unilocular, singly ostiolate, upto430µmwideand190–300µmtall,includingshortneckswhichrarelyoccur.Pycnidialwallconsistingoutoftworegionsof textura angularis;theouterregionbrown,2–3cellsthick,5–7µmwide,innerregionbrown,3–4cellsthick,7–15µmwide,withtheoutsidecellscompressed.Conidial mass globose or in cirrhi, white, pale-yellow to yellow, but predominantly pale-yellow.Alpha conidiophores cylindrical, somefiliform, rarelyseptateandbranched,5–35×1–3µm(av.=25×2µm).Alpha conidiogenous cells subcylindrical, tapering towards the apex, collarettesandpericlinalthickeningpresent,3–19× 1–2 µm (av.=10×1.5µm).Alpha conidia commonly found, fusoid-ellipsoidal, apex acutely rounded, base obtuse to subtruncate, multi-guttulate with guttules grouped at the polar ends, rarely biguttulate,(7–)9.5–10.5(–13)×(1.5–)2–3(–3.5)µm(av.=10×2.5µm).Beta conidiophores ampulliform to subcylindrical, rarely branched, 10–34 ×1–2µm(av.=26×1.5µm).Beta conidiogenous cells subcylindrical, tapering towards the apex, collaretteandpericlinalthickeningpresent,7–14×1–2µm(av.=11–1.5µm).Beta conidia less common than alpha conidia, straight,curvedorhamate,20–25×0.5–1µm(av.=23–1µm).Gamma conidia rarely observed, fusoid to subcylindrical, apex acutely rounded,basesubtruncate,multi-guttulate,12–18× 1.5–2µm(av.=15×2µm).DescriptionadaptedfromMostertetal.(2001a).
Specimens examined.frAnCe, Bordeaux,Naujan-et-Postiac, onVitis vinifera (CabernetSauvignongrapevine),May1998,P. Larignon (PREM56460neotype,ex-neotypecultureCBS114016).–itAly, Perugia, on Vitis vinifera,May1980,A. Zazzerini(CBS267.80=CPC2671).–turkey, from Vitis vinifera,1Dec.2001,M. Erkan(CBS114867=CPC4708).–USA, California, on Vitis vinifera, J.D. Cucuzza (CBS111888=ATCC48153=CPC2673).
Notes—Grove(1919)distinguishedP. ampelina(K58408)from P. viticolabyitsexternalappearanceonthehost.How-ever,Mostertetal.(2001a)re-examinedthetypespecimen,andfoundalphaconidiatobeellipsoid-fusoid,8–12×2.5–3.5µm, within the range of P. viticola(Mostertetal.2001a:f.29),andthusconsideredthemtobesynonymous.Udayangaetal.(2012)proposedD. neoviticolaasanom.nov.forP. viticola, but this name is superfluous, as the older epithet ‘ampelina’hasprecedenceandshouldbeadopted.Diaporthe ampelina (clade53) isawell-resolvedspecies. Itcauses cane and leaf spot and infections of pruning wounds of Vitis and Ampelopsidisspp.(Vitaceae).SeveralspeciesofDiaporthe can infect the host and cause variable symptoms in differentpartsof thevine(canes, leavesand fruits)causingconsiderable confusion in the taxonomy of these species on grapevine(Phillips1999,Scheperetal.2000,Mostertetal.
Fig. 2 Diaporthe anacardii (CBS720.97).a.ConidiomatasporulatingonPNA;b.conidiomatasporulatingonPDA;c,d.conidiogenouscells;e.betaconidia;f.alphaconidia.—Scalebars=10µm.
15R.R.Gomesetal.:Diaporthe
2001a).Merrinetal.(1995)studiedthevariationofDiaporthe in Australiausingmorphology.Theyidentifiedtwotaxa(Phomopsistaxon1andtaxon2),whichcausecaneandleafblightofVitisspp.;andtaxon2wasidentifiedasshowingmoreresem-blance to P. viticola.Mostertetal.(2001a)studiedthespeciesoccurring on grapevines in South Africa using morphological, cultural, molecular and pathological characterisation and clarifiedthetaxonomyofthiscomplex.Diaporthe ampelina(=Phomopsis viticola, D. neoviticola, Phomopsis taxon 2 from Australia)was found tobe thecauseofcaneand leafspotdisease,andwasneotypified.Althoughthesexualmorphhasneverbeenreported,Santosetal.(2010)foundbothMATlocitobepresentinthisspecies,andshowedthatitisheterothallic.However,thesexualmorphcouldnotbeinducedinculturebycrossingopposingmatingtypes.
Diaporthe amygdali(Delacr.)Udayanga,Crous&K.D.Hyde,Fung.Diversity56:166.2012
Basionym.Fusicoccum amygdaliDelacr.,Bull.Soc.Mycol.France21:280.1905. ≡Phomopsis amygdali(Delacr.)J.J.Tuset&M.T.Portilla,Canad.J.Bot.67,5:1280.1989.
Specimens examined.PortugAl,Mirandela,fromPrunus dulcis, 2010, E. Diogo(ex-epitypecultureCBS126679);Tavira,onPrunus dulcis, 2010, E. Diogo(CBS126680).–south AfriCA,WesternCapeProvince,onVitis vinifera,1Mar.1997,L. Mostert(CBS111811=CPC2632);WesternCapeProvince, in wood on Prunus salicina,2008,U. Damm (CBS120840=CPC5833).–usA,Georgia,cankersonPrunus persica,Mar.1994,W. Uddin (CBS115620=FAU1005).
Notes — Diaporthe amygdali(clade42)isthecausalagentoftwigcankerandblightofalmonds(Prunus dulcis)andpeach(P. persica)whereverthesehostsaregrown(Diogoetal.2010).ItwasfirstdescribedasFusicoccum amygdali causing cankers onalmondsinFrance(Delacroix1905).Tuset&Portilla(1989)re-examined the type specimen of F. amygdali and, based on morphology and symptomatology, they considered that it would be best accommodated in the genus Phomopsis. Clade42containstheex-epitypestrain(CBS126679),fivePhomopsis amygdali isolates from Prunus dulcis in Portugal, from P. persicainUSA,P. salicina in South Africa, and from Vitis vinifera inSouthAfrica.
Diaporthe anacardii(Early&Punith.)R.R.Gomes,C.Glienke& Crous, comb. nov.—MycoBankMB802923;Fig.2
Basionym.Phomopsis anacardiiEarly&Punith.,Trans.Brit.Mycol.Soc.59,2:345.1972.
Conidiomata pycnidial, sporulating profusely on OA, globose, up to600µmdiam,multilocular,black,erumpent;creamconidialdropletsexudingfromcentralostioles;wallsconsistingof3–6layers of medium brown textura angularis.Conidiophores hya-line, smooth, 1–3-septate, branched, densely aggregated, cy-lindrical,straighttosinuous,10–25×2–3µm.Conidio genous cells9–16×1.5–2µm,phialidic,cylindricaltocymbiform,termi-nalandlateral,withslighttapertowardsapex,1–1.5µmdiam,withvisiblepericlinalthickening;collaretteslightlyflared,upto2µmlongwhenpresent.Paraphyses rarely present, hyaline, smooth, 1–3-septate, cylindrical with obtuse ends, extending aboveconidiophores.Alpha conidia aseptate, hyaline, smooth, guttulate, fusoid to ellipsoid, tapering towards both ends, straight, apex subobtuse, base bluntly rounded with flattened hilum,(6.5–)7–8(–9)×(2–)3(–3.5)µm.Gamma conidia not observed.Beta conidia spindle-shaped, aseptate, smooth, hyaline, apex subacutely rounded, base truncate, tapering from lowerthirdtowardsapex,curved,(15–)20–25×1.5(–2)µm. Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C.OnOAdirtywhitewithmoderateaerialmyceliumandpatchesofiron-grey.OnPDAhavingpatchesofdirtywhiteandumber,reversebaywithpatchesofumber.OnMEAhavingpatchesofdirtywhiteandolivaceous-grey,reverseumberwithpatchesofolivaceous-grey.
Specimens examined.eAst AfriCA, on Anacardium occidentale,Apr.1997,M. Puccioni(epitypedesignatedhereCBSH-21101,cultureex-epitypeCBS720.97).–kenyA, on Anacardium occidentale,4Dec.1969,M.P. Early (holotypeIMI144866).
Notes — Phomopsis anacardii (clade69asD. anacardii)was described from Anacardi occidentalis in Kenya, and also recorded fromNigeria,GuineaandCuba (Early&Punithal-ingam1972).
Fig. 3 Diaporthe angelicae (CBS111591).a,b.Transversesectionthroughconidiomata,showingconidiomatalwall;c,d.conidiogenouscells;e.alphaandbetaconidia;f.conidiogenouscellsgivingrisetobetaconidia;g.betaconidia.—Scalebars:a=140µm,allothers=10µm.
16 Persoonia–Volume31,2013
Diaporthe angelicae(Berk.)D.F.Farr&Castl.,Mycoscience44:204.2003.—Fig.3
Basionym.Sphaeria angelicaeBerk.,Mag.Zool.Bot.:28.1837. ≡ Diaporthopsis angelicae(Berk.)Wehm.,ThegenusDiaportheNitschke:228.1933. ≡ Mazzantia angelicae(Berk.)Lar.N.Vassiljeva,PyrenomycetesoftheRussiaFarEast.I.Gnomoniaceae:49.1993. =Leptosphaeria nigrellaAuersw.,Mycol.Eur.Pyr.5/6,pl.12,f.163.1869. ≡ Diaporthe nigrella(Auersw.)Niessl,Beitr.:51.1872. ≡ Diaporthopsis nigrella(Auersw.)Fabre,Ann.Sci.Nat.,Bot.615:35.1883.
Conidiomata pycnidial, globose to ellipsoidal, aggregated or scattered,darkbrowntoblack,immersed,ostiolate,100–281μmwide,70–200μmtall, lackingnecks,withoutersurface covered inhyphae; pycnidal wall consisting of brown, thick-walled cells of textura angularis;conidialmassgloboseorexud-ingincirrhi,whitetopaleluteousorpaleyellow.Conidio phores hyaline, subcylindrical, rarely branched, tapering towards the apex,aseptate,(12–)13–16(–18)×3(–4)μm.Conidiogenous cells hyaline, subcylindrical, straight to curved, tapering towards the apex, collarette not flared, periclinal thickening inconspicu-ous,8–10(–11)×3(–3.5)μm.Alpha conidia hyaline, oblong to ellipsoid, apex bluntly rounded, base obtuse to subtruncate, bi-tomulti-guttulate,(7–)8–10(–11)×3(–4)μm.Beta conidia hyaline,smooth,spindleshaped,slightlycurved,(19–)22–26 (–28)×(1–)2μm.Gamma conidia notobserved(CBS111592). Culture characteristics — See Castlebury et al.(2003).
Specimens examined.AustriA, Karnten, St.Margareten, decayingstems of Heracleum sphondylium,Aug.2001,A.Y. Rossman(CBS111591=AR3724);Niederosterreich,Ottenstein, decaying stems of Heracleum sphondylium,Aug.2001,A.Y. Rossman(ex-epitypecultureCBS111592=AR3776).–frAnCe,Bretagne,LaVilleBorée,nearQuessoy,onseedsofHeracleum sphondylium, 27July1990,H.A. van der Aa(CBS501.90);seadunes near Seignose le Penon, on Eryngium maritimum, leafspots,10June1986,H.A. van der Aa(CBS344.86).–itAly, SanCasciano,Prov.,Florence,twig blight of Foeniculum vulgare,July1996,L. Mugnai(CBS100871).–PortugAl, MalveiradaSerra,Sintra,onFoeniculum vulgare, A.J.L. Phillips (CBS123215 =Ph-C133/1).
Notes — Diaporthe angelicae(clade17)isknowntocausestem decay in several hosts including Heracleum sphondylium (Apiaceae)and Foeniculum vulgare(Apiaceae)inEuropeandNorthAmerica(Santos&Phillips2009).Wehmeyer(1933)not only linked the conidial form of Phomopsis asteriscus to the sex- ual state Diaporthopsis angelicae, but also stated that Diaporthe berkeleyi was a synonym of Diaporthopsis angelicae.However,Castleburyetal.(2003)showedthatDiaporthopsis is a synonym of Diaporthe, and also designated an epitype for D.angelicae.
Diaporthe arctii (Lasch)Nitschke,PyrenomycetesGermanici2:268.1870
Basionym.Sphaeria arctii Lasch, inRabenh.,Klotzsch.Herb.VivumMycol.:no.1046.1846. ≡ Phomopsis arctii (Lasch)Traverso,Fl.Ital.Crypt.,Pars1:Fungi.Pyre-nomycetae.Xylariaceae,Valsaceae,Ceratostomataceae:226.1906.
Specimen examined.unknown, from Arctium sp., Sept.1925,A.W. Archer (CBS136.25).
Notes—There are several clades that contain isolatespreviously identified asD. arctii (clades 16, 19, part 2 and67,part4).Wesuspectthatclade19mayrepresentthereal D. arctii, as it is basal to D. subordinaria,andWehmeyer(1933)regardedthelatter(fromPlantago lanceolata)assynonymofD. arctii(fromArctium).
Diaporthe arecae (H.C.Srivast.,Zakia&Govindar.)R.R. Gomes,C.Glienke&Crous,comb. nov.—MycoBank MB802924
Basionym.Subramanella arecaeH.C.Srivast.,Zakia&Govindar.,Myco-logia54,1:7.1962.
Specimens examined.indiA, on fruit of Areca catechu,Feb.1964,H.C. Srivastava(isotypeCBSH-7808,ex-isotypecultureCBS161.64).–surinAme, on fruits of Citrussp.,Oct.1975,I. Block(CBS535.75).
Notes—TheDiaporthe isolate fromcitrus (CBS535.75)couldwellbedistinct,butmorestrainsarerequiredtoresolvethisclade(clade87).
Diaporthe arengae R.R.Gomes,C.Glienke&Crous,sp. nov. —MycoBankMB802925;Fig.4
Etymology.Namedafter thehost genus fromwhich itwas collected,Arenga.
Pycnidia in culture on PNA sporulating poorly, subglobose, up to 250µmdiam,black,erumpent;creamconidialdropletsexudingfromcentralostiole;wallsconsistingof3–6layersofmediumbrown textura angularis.Conidiophores hyaline in upper region, palebrownatbase,smooth,0–6-septate,branched,denselyaggregated,cylindrical,straighttosinuous,10–60×2.5–4µm.Conidiogenous cells8–15×1.5–2.5µm,phialidic,cylindrical,terminalandlateral,withslighttapertowardsapex,1–1.5µmdiam,withvisiblepericlinalthickening;collarettenotflared,upto2µmlongwhenpresent.Paraphysesnotobserved.Alpha conidia aseptate, hyaline, guttulate, fusoid-ellipsoid, tapering towards both ends, apex subobtuse, base with flattened hilum, (5–)6–7(–9)×(2–)2.5(–3)µm.Gamma conidianotobserved.Beta conidia rarely observed, subcylindrical, aseptate, smooth, hyaline, apex bluntly rounded, base truncate, tapering absent toveryslight,curved,20–25×1.5µm.
Fig. 4 Diaporthe arengae (CBS114979).a.ConidiomatasporulatingonPNA;b,c.conidiogenouscells;d.betaconidia;e,f.alphaconidia.—Scalebars=10µm.
17R.R.Gomesetal.:Diaporthe
Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C.OnMEAsurfacewithfluffyaerialmycelium,paleluteous,inreverseorangewithpatchesofsienna.OnOAumber with patches of sienna and saffron, in reverse umber withpatchesofsaffron.OnPDAsurfacewithfluffywhiteaerialmycelium, umber with patches of saffron, in reverse umber with patchesofpaleluteoustoluteous.
Specimen examined.hong kong,VictoriaPeak,fromArenga engleri,7Oct.1999,K.D. Hyde(holotypeCBSH-21104,cultureex-typeCBS114979=HKUCC5527).
Notes—TheDiaporthe species occurring on palms are sum-marisedbyFröhlichetal.(1997).Diaporthe arengae(clade81)is distinguished from known species based on a combination ofitsconidialmorphologyandhost.
Diaporthe aspalathiE.Jansen,Castl.&Crous,Stud.Mycol.55:71.2006
Basionym.Diaporthe phaseolorum var.meridionalis F.A. Fernández,Mycologia88: 438. 1996 (nonD. meridionalisSacc.,Syll. Fung.1: 638.1878).
Specimens examined.south AfriCA,WesternCapeProvince,Clanwilliam,Langebergpunt,inbranchonAspalathus linearis, J.C. Janse van Rensburg (ex-typecultureCBS117169=CPC5428);incrownonAspalathus linearis, 17Oct.1997,J.C. Janse van Rensburg (CBS117168=CPC5420);onAspalathus linearis,2Dec.1996, S. Lamprecht(CBS117500=CPC5408).
Notes — Diaporthe aspalathi (clade 93) causes soybeanstemcankerintheSouth-easternUSA(Fernández&Hanlin1996),andisnotcloselyrelatedtoD. phaseolorum as might beexpected.Althoughmorphologically similar, this speciesclustered apart from the reference strain of D. phaseolorum (clade4).Diaporthe aspalathi is also the main causal organism ofcankeranddiebackofrooibos(Aspalathus linearis),andnotD. phaseolorum asreportedearlier(Smit&Knox-Davies1989a,b,vanRensburgetal.2006).
Diaporthe australafricanaCrous&VanNiekerk,Australas.Pl.Pathol.34:33.2005
Specimens examined.AustrAliA, on Vitis vinifera,1July1995,R.W.A. Schepers(ex-typecultureCBS111886=CPC2676).–south AfriCA, on V. vinifera,1Nov.1997,L. Mostert(CBS113487=CPC2655).
Notes—Clade49contains two isolatesofD. australafricana,oneofthembeingtheex-typestrain(CBS 111886),whichis a sibling species of D. viticola inclade50(vanNiekerketal.2005). Both species were described from Vitis vinifera, but D. australafricana is thus far only known from grapevines in AustraliaandSouthAfrica.
Diaporthe batatas Harter&E.C.Field,Phytopathology2:121. 1912
Specimen examined.USA,on Ipomoea batatas, Feb.1921, L.L. Harter (CBS122.21).
Notes—Clade8consistsofasinglestrainofD. batatas isolated from Ipomoea batatasintheUSA.ThisspeciesandD. phaseolorum have in the past been considered as varieties, namely D. phaseolorumvar.batatatis and D. phaseolorumvar.batatae.However,thegeneticdatarevealednohomologybe-tweenthetwospecies.AlthoughitisnotcertainifCBS122.21(culturesterile)isanex-typestrainofD. batatas, it is regarded asauthenticforthename.
Diaporthe beckhausiiNitschke,PyrenomycetesGermanici2:295.1870
Specimen examined.unknown, from Viburnum sp.,Sept. 1927,L.E. Wehmeyer(CBS138.27).
Notes—Clade47isrepresentedbyD. beckhausii, which was isolated from Viburnum sp.(originunknown,presumablyNorth America, whereas the species was originally described from Viburnumcollected inGermany).Diaporthe beckhausii is known from woody stems of Betulasp.,Cydonia japonica, Elaeagnus angustifolia, Halesiasp., Menispermum canadense, Menispermum sp.,V. opulus, Viburnum sp. and V. tinus in temperateNorthAmericaandEurope(Farr&Rossman2012).
Diaporthe brasiliensis R.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802926;Fig.5
Etymology.Namedafterthecountrywhereitwascollected,Brazil.
Conidiomata pycnidial, globose to conical, immersed, scattered or aggregated, brown to black, ostiolate, 70–160μmwide,
Fig. 5 Diaporthe brasiliensis (CBS133183).a.ConidiomatasporulatingonPDA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall; d,e.conidiogenouscells;f,g.alphaconidia.—Scalebars:b=80µm,allothers=10µm.
18 Persoonia–Volume31,2013
60–140μmtall,necks60–130μmtall,outersurfacesmooth;pycnidal wall consisting of brown, thick-walled cells of textura angularis;conidialmassglobose,whitetopale-luteous.Conidio phoreshyaline,cylindrical,filiform,straighttocurved,1–3-sep-tate,(17–)20–27(–30)×2(–4)μm.Alpha conidiogenous cells hyaline,cylindrical,filiform,straighttocurved,collaretteflared,withslightpericlinalthickening,(7–)8–12(–14)×2(–3)μm.Al-pha conidia hyaline, ellipsoid to irregular, apex bluntly rounded, baseobtusetosubtruncate,bi-tomulti-guttulate,6–7(–8)× 2–3μm.Beta and gamma conidianotobserved. Culturecharacteristics—ColoniesonPDAflat,withanentireedge, surface mycelium dense and felty, buff, grey-olivaceous orolivaceous-grey;coloniescoveringdishafter2wkat25°Cinthedark;reverseolivaceous,dullgreen,olivaceous-buff.OnOA raised, entire edge, surface mycelium dense felty, smoke-greytogrey-olivaceous;reversepurplishgreytopalepurplishgrey,greyolivaceousorolivaceousbuff.OnMEAraised,withan entire edge, buff, smoke-grey, with patches of olivaceous-greyandvinaceous-buff;reversedarkmouse-grey,buff.
Specimens examined. BrAzil, RiodeJaneiro, endophytic species isolated from leaf of Aspidosperma tomentosum(popularnamePeroba-do-campo), July 2007, K. Rodriguez (holotypeCBSH-21100, ex-type cultureCBS133183=LGMF924=CPC20300);samecollectiondetails(LGMF926=CPC20302).
Notes—Endophytic isolates (clade36) fromamedicinalplantinBrazil.
Diaporthe carpini(Pers.)Fuckel,Jahrb.NassauischenVereinsNaturk.23–24:205.1870(1869–1870)
Basionym.Sphaeria carpiniPers.,Syn.Meth.Fung.(Göttingen)1:39.1801.
Specimen examined.sweden,Skåne,S.Mellbypar.,Stenshuvud,onCarpinus betulus,14Apr.1989,K. & L. Holm(CBS114437=UPSC2980).
Notes — Diaporthe carpini (clade55) is known from several European countries, where it occurs on Carpinus spp.
Diaporthe caulivora (Athow&Caldwell)J.M.Santos,Vran-dečić&A.J.L.Phillips,Persoonia27:13.2011
Basionym.Diaporthe phaseolorum var. caulivora Athow & Caldwell, Phytopathology44:323.1954.
Specimens examined.CAnAdA, Ontario, in mature stem on Glycine soja, Mar.1955,A.A. Hildebrand (CBS178.55=ATCC12048=CECT2023).– CroAtiA, in stem on Glycine max,K.Vrandečić(ex-neotypecultureCBS127268).
Notes—Clade91isrepresentedbytwoisolatesofD. caulivora on Glycine soja and G. max, respectively obtained from Canada(CBS178.55)andCroatia(ex-neotype:CBS127268).ThesoybeancankerspeciescomplexwasrecentlytreatedbySantosetal.(2011).
Diaporthe celastrinaEllis&Barthol.,J.Mycol.8,4:173.1902 Specimen examined.unknown, on Celastrus scandens,Sept.1927,L.E. Wehmeyer(CBS139.27).
Notes—StrainsfromtheUSAarerequiredtoconfirmtheidentityofthisculture(clade66).
Diaporthe chamaeropis(Cooke)R.R.Gomes,C.Glienke&Crous, comb. nov.—MycoBankMB802927;Fig.6
Basionym.Phoma chamaeropisCooke,Grevillea13(no.68):95.1885. ≡ Phomopsis chamaeropsis(Cooke)Petr.,as‘Phomopsis chamaeropis’,Ann.Mycol.17,2/6:83.1920(1919).
Conidiomata pycnidial in culture on PNA, globose, up to 400 µm diam(upto600µmdiamonOA),black,erumpent;creamco-nidialdropletsexudingfromcentralostioles;wallsconsistingof3–6layersofmediumbrowntextura angularis.Conidio phores hyaline,smooth,1–5-septate,branched,denselyaggregated,cylindrical,straighttosinuous,10–50×2–2.5µm.Conidiogenous cells 10–20 ×1.5–2µm,phialidic,cylindrical,terminalandlateral,withslighttapertowardsapex,1–1.5µmdiam,withvisiblepericlinalthickening;collarettenotobserved.Paraphyses notobserved.Alpha conidia aseptate, hyaline, smooth, gut-tulate, fusoid to ellipsoid, tapering towards both ends, straight, apex subobtuse, base subtruncate, (5–)6–8(–9)× 2(–2.5)µm.Gamma conidia not observed.Beta conidia spindle- shaped, aseptate, smooth, hyaline, apex acutely rounded, base truncate, tapering from lower third towards apex, curved, (20–)22–27(–30)×1.5(–2)µm. Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C.OnOAwithmoderateaerialmycelium,surface dirty white with patches of pale olivaceous-grey, re-versewithpatchesofdirtywhiteandsienna.OnMEAsurface
Fig. 6 Diaporthe chamaeropis (CBS454.81).a.ConidiomatasporulatingonPDA;b.conidiomatasporulatingonPNA;c–e.conidiogenouscells;f.alphaconidia;g.betaconidia.—Scalebars=10µm.
19R.R.Gomesetal.:Diaporthe
dirty white with patches of olivaceous-grey, reverse sienna, withpatchesofluteous.OnPDAsurfaceolivaceous-greywithpatchesofdirtywhite,reverseiron-grey.
Specimens examined.CroAtiA,Rab,slopebehindHotel ‘Imperial’,ondead branch of Spartium junceum,July1970,J.A. von Arx(CBS753.70).–greeCe,Thessaloniki,deadpartofleafof Chamaerops humilis,Aug.1981,H.A. van der Aa(CBS454.81).
Notes—Conidialdimensionscloselyfitthoseprovidedintheoriginaldescription(onChamaerops humulisfromCzecho-slovakia;Uecker1988),suggestingthatthesecultures(clade77)couldbeauthenticforthename.
Diaporthe cinerascens Sacc.,Syll.Fung.(Abellini)1:679.1882. —Fig.7
=Phoma cinerescensSacc.,Michelia1(no.5):521.1879. ≡ Phomopsis cinerascens(Sacc.)Traverso,Fl.Ital.Crypt.Pyrenomycetae2,1:278.1906.
Conidiomata pycnidial,sporulatingpoorlyonMEA,globose,upto300µmdiam,black,erumpent;creamy-luteousconidialdropletsexudingfromcentralostioles;wallsconsistingof3–6layers of medium brown textura angularis.Conidiophores hyaline, smooth, 1–3-septate, branched, densely aggregated, cylindrical,straighttosinuous,17–30×2–3µm.Conidiogenous cells8–18× 2–3 µm, phialidic, cylindrical, terminal and lateral, withslight taper towardsapex,1.5–2µmdiam,withvisiblepericlinal thickening; collarettemostly absent, slightly flaredwhenpresent,upto2µmlong.Paraphysesnotobserved.Alpha conidia aseptate, hyaline, smooth, guttulate, fusoid to ellipsoid, tapering towards both ends, straight, apex subobtuse, base subtruncate,7–8(–9)×(2.5–)3µm.Gamma conidia aseptate, hyaline, smooth, ellipsoid-fusoid, apex acutely rounded, base subtruncate,8–12×3µm.Beta conidianotobserved. Culture characteristics — Colonies covering dish after 2 wk in thedarkat25°C.OnMEAwithprofuseaerialmycelium,surfacedirtywhite,reverseochreouswithpatchesofumber.OnPDAwithsparseaerialmycelium,surfaceolivaceous-grey,reverseiron-grey.OnOAsurfacewithmoderateaerialmyce-lium,olivaceous-greytopaleolivaceous-grey.
Specimen examined.BulgAriA,Kostinbrod,PlantProtectionInstitute,onbranch of Ficus carica,1995,E. Ilieva(CBS719.96).
Notes — Diaporthe cinerascens (clade 76) represents aEuropean species occurring on Ficus, so the present culture could be authentic for the name, as the conidial dimenions matchthoseprovidedintheoriginaldescription.Thisspecieswas orginally associated with canker and dieback of Ficusspp.inItaly(Saccardo1879),andthecausalorganismidentifiedasPhomopsis cinerascens(sexualmorph:Diaporthe cinerascens)byGrove(1935).Diaporthe cinerascens affects all commercial figs inCalifornia (Ogawa&English 1991), and is found inseveralgeographicallocationsoftheworld(Hampson1981,
Anderson&Hartman1983,Benschopetal.1984,Banihashemi&Javadi2009).Ficusspp.areimportantexoticgardenorna-mentalsacrosstheUSAandCanadaaswellasinthetropics.
Diaporthe citri F.A.Wolf,J.Agric.Res.33,7:625.1926
=Phomopsis citriH.S.Fawc.,Phytopathology2,3:109.1912.
Specimens examined.BrAzil, on seed of Glycine max, A. AlmeidaEM-BRAPA/PR(LGMF946=CPC20322).–itAly, unknownhost,June1939,G. Goidánich(CBS199.39).–surinAme, Paramaribo, on decaying fruit of Citrus sinensis,Apr.1932,N.J. van Suchtelen(CBS230.52).
Notes—Clade6isrepresentedbythreeisolates.Oneiso-late(CBS199.39)waspreviouslyidentifiedasD. conorum from Italy,whileanotheroriginatesfromsoybeanseedcollectedinBrazil(LGMF946),andthethirdisolateisfromCitrus sinensis inSuriname(CBS230.52).BecauseD. conorum is regarded as synonym of D. eres (clade67),wetentativelyrefertothiscladeas D. citri, awaiting more isolates from Citrus.Diaporthe citri is a serious pathogen that is widely distributed, and associated withmelanosisandstem-endrotofcitrusfruits(Punithalingam&Holliday1973,McKenzie1992,Mondaletal.2007,Farr&Rossman2012).
Diaporthe convolvuli (Ormeno-Nuñez,Reeleder&A.K.Wat- son)R.R.Gomes,C.Glienke&Crous,comb. nov.—Myco-BankMB802928
Basionym.Phomopsis convolvuliOrmeno-Nuñez,Reeleder&A.K.Wat-son, Canad.J.Bot.66,11:2232.1988.
Specimen examined. turkey, isolated from leaves with anthracnose on Convolvulus arvensis, D. Berner (CBS124654=DP0727).
Notes — Phomopsis convolvuli (clade 3) was originally described from diseased leaves of Convolvulus arvensis in Québec(Ormeno-Nuñezetal.1988).TheisolateofPhomopsis convolvulistudiedhere(CBS124654),wasfoundcausinganthracnoseonfieldbindweed(Convolvulus arvensis),atrou-blesome perennial weed to many important agricultural crops in the world, and was considered potentially useful as biological controlagent(Kulecietal.2009).
Diaporthe crataegi(Curr.)Fuckel,Jahrb.NassauischenVer-einsNaturk.23–24:204.1870
Basionym.Valsa crataegi Curr.,Trans.Linn.Soc.London22:278.1858.
Specimen examined.sweden,Skåne,Trolle-Ljungbypar.,Tosteberga,onCrataegus oxyacantha,15Apr.1989,K. & L. Holm(CBS114435=UPSC2938).
Notes — Clade 41 is represented by D. crataegi isolated from Crataegus oxyacantha inSweden.Thespeciesiscommonon C. chrysocarpa, C. laevigata and C. oxyacantha in Canada andEurope(Farr&Rossman2012).
Fig. 7 Diaporthe cinerascens (CBS719.96).a.ConidiomatasporulatingonPDA;b,c.conidiogenouscells;d.alphaconidia.—Scalebars=10µm.
20 Persoonia–Volume31,2013
Diaporthe crotalariaeG.F.Weber,Phytopathology23:602. 1933
=Phomopsis crotalariaeG.F.Weber,Phytopathology23:602.1933.
Specimen examined.USA, onCrotalaria spectabilis,Oct. 1933,G.F. Weber(ex-typecultureCBS162.33).
Notes—Clade92containstheex-typestrain(CBS162.33)of D. crotalariae isolated from Crotalaria spectabilisintheUSA.
Diaporthe cuppatea(E.Jansen,Lampr.&Crous)Udayanga,Crous&K.D.Hyde,Fung.Diversity56:166.2012
Basionym.Phomopsis cuppateaE.Jansen,Lampr.&Crous,Stud.Mycol.55:72.2006.
Specimen examined. south AfriCA,WesternCapeProvince,onAspalathus linearis,2006,J. Janse van Rensburg(holotypeCBSH-19687,ex-typecultureCBS117499=STE-U5431=CPC5431).
Notes — Diaporthe cuppatea(clade20)isknownonlyfromthe original collection made from dying branches of Aspalathus linearisinSouthAfrica(vanRensburgetal.2006).
Diaporthe cynaroidis Marinc.,M.J.Wingf.&Crous,CBSBio-diversitySer.(Utrecht)7:39.2008
Specimen examined.south AfriCA,WesternCapeProvince,onleaflitterof Protea cynaroides,26June2000,S. Marincowitz(ex-typecultureCBS122676=CMW22190=CPC13180).
Notes—Clade48containstheex-typecultureofD. cynaroidis(CBS122676),whichwas isolated from Protea cynaroides inSouthAfrica(Marincowitzetal.2008).Thisspeciesiscloselyrelated to D. australafricana and D. viticola (clades49and50,respectively).
Diaporthe decedens (Pers.)Fuckel,Jahrb.NassauischenVer- einsNaturk.23–24:30.1871
Basionym.Sphaeria tessella var.decedens Pers., Syn.Meth. Fung.(Göttingen)1:48.1801.
Specimens examined.AustriA, on Corylus avellana,Oct.2001,W. Jaklitsch(CBS109772=AR3459).–sweden,Öland,Kastlösapar.,onCorylus avellana,7June1989,K. & L. Holm(CBS114281=UPSC2957).
Notes — Diaporthe decedens represents a European spe-cies on Corylus.Clade68consistsoftwoisolatesobtainedonCorylus avellanafromAustriaandSweden.
Diaporthe detrusa (Fr.)Fuckel,Jahrb.NassauischenVereinsNaturk.23–24:205.1870(1869–1870)
Basionym.Sphaeria detrusaFr.,inKunze&Schmidt,MykologischeHefte(Leipzig)2:43.1823.
=Phoma detrusaSacc.,Michelia2:96.1880. ≡ Phomopsis detrusa(Sacc.)Traverso,Fl.Ital.Crypt.Pars1:Fungi.Py-renomycetae.Xylariaceae,Valsaceae,Ceratostomataceae1,1:195.1906.
Specimens examined.AustriA, on Berberis vulgaris,Oct. 2001,A.Y. Rossman(CBS109770=AR3424).–sweden,Uppland,Hållnäspar.,onBerberis vulgaris,14May1991,K. & L. Holm(CBS114652=UPSC3371).– unknown, on Berberis vulgaris,Sept.1927,L.E. Wehmeyer(CBS140.27).
Notes—Clade 54 contains three isolates of D. detrusa obtained from Berberis vulgaris in Austria, Sweden and one ofthemwithanunknownorigin(presumablyNorthAmerica).ThisEuropeanspeciesisknowntoalsooccurintheUSA(Farr&Rossman2012).
Diaporthe elaeagni Rehm,Syll.Fung.14:546.1899.—Fig.8
?=Phoma elaeagniSacc.,Michelia1,3:354.1878. ≡ Phomopsis elaeagni(Sacc.)Petr.,Ann.Mycol.19,1–2:48.1921.
Specimen examined.netherlAnds,Maassluis,ontwigofElaeagnussp.,May1972,J. Gremmen(CBS504.72).
Notes—IncultureCBS504.72(clade73)primarilyproducesbetaconidia(spindleshaped,16–22× 2 µm, thus wider than seenonaverageinmostotherspecies);alphaconidiararelyobserved,fusoid-ellipsoidal,7–10× 2–3 µm, thus correlating with dimensions of Phomopsis elaeagni (Sacc.)Petr.,whichis a homonym of P. elaeagniSacc.Furthermore,conidialdi-mensions of the asexual state of D. elaeagniarenotknown.Additional collections and type studies are thus required toresolve the complex occurring on Elaeagnus.
Diaporthe endophyticaR.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802929
Etymology. Named after its endophytic growth habit.
Culturessterile.Diaporthe endophytica(clade5)differsfromits closest phylogenetic neighbour, D. phaseolorum(clade4),byuniquefixedallelesinfivelocibasedonalignmentsoftheseparate loci deposited inTreeBase as studyS13943: ITSpositions357(C),359(G),360(T),368(A),369(A),371(A),372(G)and373(G);TUBpositions135(C)and592(T);CALposition145(G);TEF1positions18(G),26(T),40(T),42(A),63(A),124(A),175(A)and343(A);HISposition369(C).
Culture characteristics — Colonies with sparse aerial my-celium,covering thedishafter2wkat25°C.OnPDAbuff,honeytoisabelline;reversesmoke-grey.OnOAsmoke-greytoolivaceous-grey.OnMEAbuffwithumberpatches;reversedarkmouse-grey,withpatchesofisabelline.
Specimens examined.BrAzil, endophytic in leaf on Schinus terebinthifolius, July2007,J. Lima(LGMF911=CPC20287,LGMF919=CPC20295),(holotypeCBSH-21107,cultureex-typeLGMF916=CPC20292
Fig. 8 Diaporthe elaeagni (CBS504.72).a.ConidiomatasporulatingonPNA;b.conidiomatasporulatingonPDA;c,d.conidiogenouscells;e.betaconidia.—Scalebars=10µm.
21R.R.Gomesetal.:Diaporthe
=CBS133811);endophyticinpetioleonMaytenus ilicifolia,July2007, R.R. Gomes(LGMF928=CPC20304, LGMF934=CPC20310,LGMF935=CPC20311,LGMF937=CPC20313); in seed on Glycine max, A. Almeida EMBRAPA/PR(LGMF948=CPC20324).
Notes—Clade5representsadistinctlineage,containingeightsterileisolatesoriginatingfromBrazil.Fourofthemwereisolated from Maytenus ilicifolia, three from S. terebinthifolius andonefromsoybeanseeds.Isolatescouldnotbeinducedtosporulateonanyofthemediadefinedinthisstudy,noronsterilisedplanthosttissueplacedonWA.
Diaporthe eres Nitschke,PyrenomycetesGermanici2:245.1870
= Phomopsis cotoneastri Punith.,Trans.Brit.Mycol.Soc.60,1:157.1973. = Phoma oblongaDesm.,Ann.Nat.Sci.Bot.20:218.1853. ≡ Phomopsis oblonga (Desm.)Traverso,Fl.Ital.Crypt.Pars1:Fungi.Pyrenomycetae.Xylariaceae,Valsaceae,Ceratostomataceae:248.1906.
Specimens examined.AustriA, on Acer campestre,Oct.2001,W. Jaklitsch (CBS109767=AR3538=WJ1643).–germAny,Monheim,onleafspotofHordeum sp.,5Aug.1984,M. Hossfeld(CBS841.84).–itAly,Milano,ontwig of Juglans regia,Dec.1980,M. Bisiach(CBS102.81).–lAtviA, on Rhododendron sp., I. Apine(CBS129168).–netherlAnds, Oostvoorne, on dead stems of Arctium sp.,13Dec.1984,M. de Nooij(CBS110.85);Soest,Dal-weg, on fallen fruit of Fraxinussp.,21Feb.1999,G. Verkley(CBS101742);Veldhoven,ondeadbranchofSorbus aucuparia,Nov.1973,W.M. Loerakker (CBS287.74);Baarn,gardenChopinlaan,ondeadbranchofWisteria sinensis,6June1983,H.A. van der Aa(CBS528.83);Soest,insidehouse,onAbutilonsp.,26Mar.1997,A. Aptroot(CBS688.97);Baarn,pottedplant,oncladodes of Opuntiasp.,23Sept.1996,H.A. van der Aa(CBS365.97);onAlliaria officinalis,Feb.1962,G.H. Boerema(CBS445.62);Baarn,ondeadleaf of Ilex aquifolium,11June1967,H.A. van der Aa (CBS370.67=MUCL9931);Prov.Zuid-Holland,HuizeOud-Poelgeest,Oegstgeest,diebackofIlex aquifolium,21Nov.1994,G.J.M. Verkley(CBS694.94);Baarn,gardenEemnesserweg90,ondeadstemofRumex hydrolapathum,19Mar.1996,H.A. van der Aa (CBS485.96);Baarn,Cantonspark,onwithering leafofMagnolia × soulangeana,23Oct.1968,H.A. van der Aa(CBS791.68);Zuid-Holland,Ridderkerk,HuystenDonck,onleaftipofOsmanthus aquifolium, 7May1977,H.A. van der Aa(CBS297.77);onPhaseolus vulgaris,Sept.1950,Goossens(CBS422.50);Baarn,ondeadstemofAllium giganteum, May1985,H.A. van der Aa(CBS283.85);fromLaburnum × watereri ‘Vossii’,Apr.1935,I. de Boer(CBS267.55);Boskoop,nursery,dyingtwigsofSkimmia japonica,Nov.1981,H.A. v. Kesteren(CBS122.82).–uk, Scotland, on living and dead twig of Fraxinus excelsior,Feb.1938,J.A. MacDonald
(CBS 250.38); onCotoneaster sp., 1971,H. Butin (ex-type culture of P. crotoneasterCBS439.82=BBAP-407=IMI162181a);Oxford,onPicea abiesseedling,Nov.1937,T.R. Peace(CBS186.37).–unknown, on rotten fruit of Malus sylvestris,May1961,Geigy(CBS375.61);May1932,W.G. Hutchinson(CBS267.32).
Notes — Diaporthe eres (clade67) is thetypespeciesofthe genus Diaporthe, and is present in several hosts, though it isknowntobemorphologicallyhighlyvariable(Castleburyetal. 2002).Wehmeyer(1933)describedthisspeciesonmorethan60hosts,andlistedseveralsynonymiesbasedonmor-phologicaldata. Adetailedmorphologicalstudyisrequiredtodesignate a suitable epitype strain for D. eres, and to resolve thestatusofallitspurportedsynonyms.
Diaporthe eugeniae (Punith.)R.R.Gomes,C.Glienke&Crous, comb. nov.—MycoBankMB802930
Basionym.Phomopsis eugeniaePunith.,Trans.Brit.Mycol.Soc.63,2:232.1974.
Specimens examined.west sumAtrA, on Eugenia aromatica,May1973,J. Waller(holotypeIMI177560);Lampung,onleafofEugenia aromatica, July1982,R. Kasim(CBS444.82).
Notes — Diaporthe eugeniae(clade83)wasoriginallyde-scribed on Eugenia aromaticafromWestSumatra.Althoughthepresent isolate could be authentic for the name, it unfortunately provedtobesterile.
Diaporthe fibrosa(Pers.)Fuckel,Jahrb.NassauischenVereinsNaturk.23–24:204.1870(1869–1870)
Basionym.Sphaeria fibrosaPers.,Syn.Meth.Fung.(Göttingen)1:40.1801.
Specimens examined.AustriA, Vienna, onRhamnus cathartica,Oct.2001, A.Y. Rossman(CBS109751=AR3425).–sweden,Uppland,Dalbypar.,Hässleborg,onRhamnus cathartica,10Mar.1987,K. & L. Holm(CBS113830=UPSC2117).
Notes—Clade52consistsoftwoisolatesfromRhamnus cathartica collectedinSwedenandAustria.Diaporthe fibrosa was originally described from Europe on Rhamnus, so these culturesmaywellprovetobeauthenticforthename.
Fig. 9 Diaporthe foeniculacea (CBS111554).a.ConidiomatasporulatingonPDA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d–f.conidiogenouscells;g.betaconidia;h.alphaconidia.—Scalebars:b=250µm,allothers=10µm.
22 Persoonia–Volume31,2013
Diaporthe foeniculacea Niessl,invonThümen,Contr.Ad.Fl.Myc.Lusit.2:30.1880.—Fig.9
=Phoma foeniculinaSacc.,Syll.Fung.3:125.1884. ≡ Phomopsis foeniculina(Sacc.)Câmara,Agron.Lusit.9:104.1947. =Phomopsis theicolaCurzi,AttiIst.Bot.Univ.Pavia,3sér.,3:65.1927. =Diaporthe neotheicolaA.J.L.Phillips&J.M.Santos,Fung.Diversity34:120.2009.
Conidiomata pycnidial, eustromatic, multilocular, immersed, ostiolate,darkbrown,scatteredoraggregated,350–890μmwide,160–320μmtall,necksabsent,outersurfacecoveredwithhyphae;pycnidalwallconsistingofbrown,thick-walledcellsoftextura angularis;conidialmassglobosetoconicalandexudingincirrhi,yellowtoreddishbrown.Conidiophores hyaline, sub-cylindricalandcylindrical,filiform,branchedabovethesepta,tapering towards the apex, 1–3-septate, (19–)20–28(–32)×2(–3)μm.Conidiogenous cells hyaline, subcylindrical and filiform,straight,slightlytaperingtowardstheapex,collarettenotflared,prominentpericlinalthickening,(10–)11–15(–17)× 2(–3)μm.Alpha conidia hyaline, oblong to ellipsoidal, apex bluntly rouded, base obtuse to subtruncate, bi- to multi-guttulate (6–)7–9× 2(–3) μm.Beta conidia hyaline, smooth, slightly curved, (26–)28–32(–34)× 1(–2) μm.Gamma conidia not observed(basedonisolateCBS111554).
Specimens examined. indiA,Calcutta,unknownhost,Feb.1948,S.R. Bose(CBS400.48).–itAly, on leaves and branches of Camellia sinensis, Oct.1927,M. Curzi(ex-typecultureofP. theicolaCBS187.27);Perugia,onDiospyros kaki,June1956,M. Ribaldi(CBS287.56);Apulia,nearBari,onPrunus amygdalus,winter1974/75,A. Ciccarone(CBS171.78).–nether-lAnds,Baarn,‘Madoera’,backfrond,onWisteria sinensis,24Apr.1969,H.A. van der Aa(CBS357.69).–new zeAlAnd,Waikatoregion,onPyrus pyrifolia, 2001, W. Kandula (CBS116957).–PortugAl,nearLisbon,SãoMarcos,base of senescent stem of Foeniculum vulgare,Apr.2002,A.J.L. Phillips (CBS111554);Évora,Foeniculum vulgare,1Nov.2007,A.J.L. Phillips(ex-type cultures of Diaporthe neotheicolaCBS123209,CBS123208);PedrasdelRei,nearTavira,onBougainvillea spectabilis,15June1988,H.A. van der Aa(CBS603.88);Madeira,SerradaAgua,baseofsenescentstemofFoeniculum vulgare,Aug.2001,A.J.L. Phillips(CBS111553).
Notes — Diaporthe foeniculacea(clade78)wasoriginallydescribed from Foeniculum vulgare in Portugal, and represents an older name for D. theicola and D. neotheicola.Therearemany described species that occur on Foeniculum vulgare(wildfennel).Amongthem,P. theicola and its teleomorph D. neotheicola (Santos&Phillips2009),and D. foeniculacea, the causal agentofstemnecrosisoffennel.Phillips(2003)redescribedD. foeniculacea, and established the sexual-asexual connec-tion between D. foeniculacea and Phomopsis foeniculina.Thesynonymy of D. neotheicola under D. foeniculacea is based on the fact that the cultures matching the original descriptions areinfactgeneticallyidentical.However,astherearenoex-type strains of D. foeniculacea, this synonymy strongly relies ontheearlieropinionofPhillips(2003).Eitherway,thismattercan only be resolved once an epitype has been designated for D. foeniculacea,fixingtheapplicationofthename.Werecom-mend that additional collections linked to stem necrosis of fennel inPortugalareobtained,beforethisdecisionismade.
Diaporthe ganjae (McPartl.)R.R.Gomes,C.Glienke&Crous,comb. nov.—MycoBankMB802932
Basionym.Phomopsis ganjae McPartl.,Mycotaxon18,2:527.1983.
Specimen examined.USA, Illinois,HannahCity,deadleaf of Cannabis sativa,depositedMar.1991,J.M. McPartland(holotypeHA10987,ex-typecultureILLS43621=CBS180.91).
Notes — Diaporthe ganjae(clade24)isknownonlyfromtheoriginal collection taken from wilted, dead leaves of Cannabis sativa in Illinois,USA (McPartland 1983). Phylogenetically D. ganjaeiscloselyrelatedtoanisolateidentifiedasD. mani
hotia(CBS505.76),isolatedfromManihot utilissimainRwanda(clade25).
Diaporthe gardeniae (Buddin&Wakef.)R.R.Gomes,C.Glienke & Crous, comb. nov.—MycoBankMB802933
Basionym.Phomopsis gardeniae Buddin&Wakef.,Gard.Chron.,ser.3103:45.1938. =Phomopsis gardeniaeH.N.Hansen&Barrett,Mycologia30,1:18.1938(homonym).
Specimen examined.itAly, on stem of Gardenia florida,June1956,M. Ri baldi(CBS288.56).
Notes — Diaporthe gardeniae (clade59)causesgardeniacanker in Gardenia jasminoides, G. lucida and Gardenia sp. (Farr&Rossman2012).Thisdiseaseisconsideredasseri-ous(Tilford1934,Huber1936,Miller1961).Itwasoriginallyobservedin1894inEngland(Cooke1894),andhassincebeenreportedfromtheUSA(Preston1945)andIndia(Mathur 1979).All parts of the plant are susceptible to infection, including roots, stemsandleaves(McKenzieetal.1940),althoughcankeredstemsarethemostdiagnosticsymptomsforthisdisease.
Diaporthe helianthi Munt.-Cvetk.,Mihaljč.&M.Petrov,NovaHedwigia34:433.1981
=Phomopsis helianthiMunt.-Cvetk.,Mihaljč.&M.Petrov,NovaHedwigia34:433.1981.
Specimens examined. serBiA, Vojvodina,overwinteringstemonHelianthus annuus, 1980,M. MuntañolaCvetkovic(ex-typecultureCBS592.81=CBSH-1540). – unknown, on seed of H. annuus,June1994,VanderhaveRes.,Rilland,Netherlands (CBS344.94).
Notes — Diaporthe helianthi(clade14)isassociatedworld-wide with stem canker and grey spot disease of sunflower (Helianthus annuus)(Muntañola-Cvetkovic´etal.1981).Yieldreductionsofupto40%havebeenrecordedinEurope(Masi-revic&Gulya1992)includingtheformerYugoslaviaaswellasFrancewhereitwasconsideredamajorpathogenofsunflower(Battilanietal.2003,Debaekeetal.2003).Diaporthe helianthi isalsowidespreadinthesunflowergrowingregionsoftheUSA(Gulyaetal.1997).Thewidegeographicdistribution,andhighgenetic variability of the pathogen lead to the evolution of new strains that could be more aggressive, causing large yield lossesandadeclineindiseasecontrol(Pecchiaetal.2004,Rekabetal.2004).
Diaporthe cf. heveae 1 Specimen examined.BrAzil,SãoPaulo, fromHevea brasiliensis,Apr.1997,D.S. Attili(CBS852.97)(originallyidentifiedasPhomopsis heveae).
Notes — Diaporthe heveae and Phomopsis heveae were both described from HeveainSriLanka,andcouldrepresentthesamespecies.TwoisolatesdepositedinCBSunderthisname,CBS852.97 (fromHevea brasiliensis in Brazil) andCBS681.84(fromHevea brasiliensis inIndia)wereshowntorepresenttwodistinctspecies(clades46and80,respectively).However,asbothwerefoundtobesterile,theirtaxonomycouldnotberesolved.
Diaporthe cf. heveae 2 Specimen examined.indiA, Kerala, Kottayam, in leaf on Hevea brasiliensis,Sept.1984,K. Jayarathnam(CBS681.84).
Notes—IsolateCBS681.84(clade80,P. heveae from Hevea brasiliensisinIndia)issterile,andthusitstaxonomycouldnotberesolved.Diaporthe heveaehasbeenreportedfromBrazil,China,India,Indonesia,Malaysia,SriLankaandThailand(Hol-liday1980,Zhuang2001,Udayangaetal.2011).
23R.R.Gomesetal.:Diaporthe
Diaporthe hickoriae Wehm.,Monogr.Gen.DiaportheNitschke&Segreg.,Univ.MichiganStud.,Sci.Ser.9:149.1933
Specimen examined.USA,Michigan,onCarya glabra,June1926,L.E. Wehmeyer (ex-typecultureCBS145.26).
Notes — Diaporthe hickoriae (clade72)occurson the bark of Carya glabraintheUSA(Wehmeyer1933).
Diaporthe hongkongensis R.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802934;Fig.10
Etymology.Namedafterthelocationwhereitwascollected,HongKong.
Conidiomatapycnidial,superficialtoembeddedonPDA,solitaryto aggregated, globose with central ostiole, exuding a creamy conidialcirrhus;pycnidialupto200µmdiam;wallof3–6layersof brown textura angularis.Conidiophores lining the inner cavity, reducedtoconidiogenouscells.Conidiogenous cells phialidic, hyaline, smooth, ampulliform to subcylindrical with prominent apicaltaper,5–12×2–4µm;apexwithpericlinalthickeningandminutecollarette,1µmlong.Paraphyses intermingled among conidiophores,hyaline,smooth,frequentlybranchedbelow,upto4-septate,withclavateterminalcell,upto80µmlong,apex2–8µmdiam.Alpha conidia hyaline, smooth, granular to gut-tulate, aseptate, fusiform, tapering towards both ends, mostly straight,apexacutelyrounded,basetruncate,(5–)6–7(–8)× (2–)2.5(–3)µm.Gamma conidia aseptate, hyaline, smooth, ellipsoid-fusoid, apex subobtuse, base truncate, 10–13 ×2µm.Beta conidia aseptate, hyaline, smooth, spindle-shaped, apex acutely rounded, base truncate, widest in mid region, mostly curvedinupperpart,18–22×1.5–2µm. Culture characteristics — Colonies covering dish after 2 wk in thedarkat25°C,withmoderateaerialmycelium.OnOAsurfacedirty white with patches of pale olivaceous-grey, reverse dirty whitewithpatchesofolivaceous-greyandiron-grey.OnPDAsurfaceiron-grey,withpatchesofdirtywhite,reverseiron-grey.OnMEAsurfacedirtywhitewithpatchesofolivaceous-grey,reverseiron-greywithpatchesofdirtywhite.
Specimen examined.hong kong,TaiPoKau,onfruitofDichroa febrifuga, 20Feb. 2002,K.D. Hyde (holotypeCBSH-21103, culture ex-typeCBS115448=HKUCC9104).
Notes—IsolateCBS115448(clade79;reportedasPhomopsis pittospori on Dichroa febrifuga fromHongKong) ismorphologically distinct from P. pittospori (fromPittosporum twigsinCalifornia;alphaconidia6–8×1.5µm,betaconidia18–20×1µm),withwideralphaandbetaconidia.
Diaporthe hordei(Punith.)R.R.Gomes,C.Glienke&Crous,comb. nov.—MycoBankMB802935
Basionym. Phomopsis hordeiPunith.,Trans.Brit.Mycol.Soc.64,3:428.1975.
Specimen examined.norwAy, Fellesbygget, As, on root of Hordeum vulgare,Oct.1992,L. Sundheim(CBS481.92).
Notes — Diaporthe hordei (clade13)wasdescribedfromHordeum vulgareintheUK.Althoughthepresentculturecouldbeauthentic(fromHordeumcollectedinNorway),itprovedtobesterile,soitsmorphologycouldnotbeconfirmed.
Diaporthe impulsa(Cooke&Peck)Sacc.,Syll.Fung.(Abel-lini)1:618.1882
Basionym.Valsa impulsa Cooke&Peck,Ann.Rep.N.Y.StateMus.Nat.Hist.27:109.1875(1874).
Specimens examined.sweden,Uppland,Dalbypar.,Jerusalem,onSorbus aucuparia,24Oct.1989,K. & L. Holm(CBS114434=UPSC3052).–unknown, on Sorbus americana,Sept.1927,L.E. Wehmeyer(CBS141.27).
Notes—Clade51isrepresentedbytwoisolatesofD. impulsa occurring on Sorbus spp. Diaporthe impulsa is a known pathogen of Sorbusspp.,andhasawidegeographicdistribu-tion(Farr&Rossman2012).ItwasoriginallydescribedfromSorbusintheUSA,thusCBS141.27maywellprovetobeagoodreferencestrainforthespecies.
Diaporthe inconspicua R.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802936
Etymology.Referringtoitsinconspicuousnature,growingasendophyteinhosttissue.
Culturessterile.Diaporthe inconspicua(clade75)differsfromitsclosestphylogeneticneighbours,clade68–74and76–78,byuniquefixedallelesinfourlocibasedonalignmentsoftheseparate locideposited inTreeBaseasstudyS13943:TUBpositions33(A),102–104and106–111(indels),127(G),149(C),151(A),195(C),204(T),357(G),446(G),449(C),465(T),484(T),559(A),592(A),629(T),653(T),708(C),732(C),754(A),763(C),784(A)and787(G);CALpositions28(C),102(G),114(T),148(T),152(T),153(A),157(C),170(G),199(C)and281(C);TEF1positions9(T),16(A),22(A),29(G),30(G),81(C),86(C),87(A),88(A),89(T),131(A),275(A),298(C)and315(T);HISpositions139(T),211(T),244(T)and408(T).
Culture characteristics — Colonies covering the dish after 2wkinthedarkat25°C.OnOAspreading,flatwithsparseaerialmycelium,surfacecreamincentre,umberinouterregion.
Fig. 10 Diaporthe hongkongensis (CBS115448).a,b.ConidiomatasporulatingonPDA;c,d.conidiogenouscells;e.betaconidia;f.alphaconidia.—Scalebars=10µm.
24 Persoonia–Volume31,2013
OnPDAsurfaceandreversecreamtodirtywhitewithsparseaerialmycelium.OnMEAwithsparseaerialmycelium,surfacebecoming folded, dirty white in centre, sienna in outer region, andluteousinreverse.
Specimens examined.BrAzil, on petiole of Maytenus ilicifolia,July2007,R.R. Gomes (holotypeCBSH-21102,ex-typecultureLGMF930=CPC20306=CBS133813);samecollectiondetails(LGMF931=CPC20307);on Spondias mombin, 2007,K. Rodriguez(LGMF922=CPC20298).
Notes—Sterileendophyticisolates(clade75)frommedici-nalplantsinBrazil.
Diaporthe infecunda R.R.Gomes,C.Glienke&Crous,sp. nov. —MycoBankMB802937
Etymology.Namedafteritssterilegrowthinculture.
Cultures sterile.Diaporthe infecunda (clade23)differs fromitsclosestphylogeneticneighbours,clade16–22,byuniquefixedallelesinfivelocibasedonalignmentsoftheseparatelocidepositedinTreeBaseasstudyS13943:ITSpositions108(T),279(C),292(G),359(C)and360(G);TUBpositions11(indel),106(G),138(T),140(A),153(G),155(T),184(A),197(G),202(C),302(A),354(A),369–374(indels),398(G),407(indel),414(C),422(T),424(G),425(C),432(G),452(C),454(C),458(C),461(G),479(T),482(T),486(C),540(T),572(C),622(A),694(T),696(T),697(G),716(C),728(C),776(G),778(G)and796(C);CALpositions64(T),83(T),104(G),146(C),151(C),155(G),159(C),172(C),176(T),179(A),184(G),197(T),206(T),212(C)and221(T);TEF1positions6(A),9(G),13(G),16(C),21(A),30(G),32(indel),39(A),40(A),41(G),42(T),43(A),79(G),83(T),90(T), 92(T),96(A),97(C),106(C),116(A),120(C),123(A),127(A),132(A),135(G),173(G),255(T),284(A),294(C),299(C)and309(A);HISpositions173(T),196(T),197(G),199(C/T),221(C),222(C),230(G),263(C),264(T),268(C),273(T)and279(C).
Culture characteristics — Colonies covering the dish after 2wkinthedarkat25°C.OnPDAsurfaceumberwithpatchesofwhite,reversechestnut.OnMEAsurfacedirtywhite,reverseumber.OnOAsurfacewithpatchesofdirtywhiteandumber.
Specimens examined.BrAzil, on leaf of Schinus terebinthifolius, July2007,J. Lima(holotypeCBSH-21095,ex-typecultureLGMF906=CPC20282=CBS133812);additionalisolateswithsamecollectiondetails(LGMF908=CPC20284,LGMF912=CPC20288,LGMF917=CPC20293,LGMF918=CPC20294,LGMF920=CPC20296); in petiole of Maytenus ilicifolia, July2007,R.R. Gomes(LGMF933=CPC20309,LGMF940=CPC20316).
Notes — Clade 23 represents endophytic isolates from leavesofmedicinalplantsgrowinginBrazil.Itconsistsofeightisolates, two from Maytenus ilicifolia, and six from Schinus terebinthifolius.
Diaporthe juglandina(Fuckel)Nitschke,PyrenomycetesGer-manici2:281.1870
Basionym.Aglaospora juglandinaFuckel,FungiRhenaniExsicc.,suppl.7(no.2101–2200):no.2159.1868.
Specimen examined.USA,Tennessee,GreatSmokyMtsNationalPark,dead wood of Juglanssp.,L. Vasilyeva(CBS121004).
Notes — Diaporthe juglandina(clade65)representsaEuro-pean taxon described from Juglans. European collections are requiredtoconfirmwhetherthisnamecanbeappliedtotheclade.
Diaporthe longispora (Wehm.)R.R.Gomes,C.Glienke& Crous, comb. nov.—MycoBankMB802938
Basionym.Diaporthe strumellavar.longispora Wehm.,Mycologia28,1:46.1936.
Specimen examined.CAnAdA,Ontario,Toronto,onRibes sp.,May1936,L.E. Wehmeyer(ex-typecultureCBS194.36).
Notes—Clade27comprisestheex-typecultureofD. stru mellavar.longispora isolated from Ribessp.,andformsasisterclade with D. sclerotioides(clade28).Diaporthe strumella is found on woody limbs, especially of Ribes spp. in temper-ateNorthAmericaandEurope (Farr&Rossman2012).As D. strumellavar.longispora is morphologically clearly a distinct species,weelevatethisvarietytospeciesstatus.
Diaporthe lusitanicae A.J.L.Phillips&J.M.Santos,Fung.Diver- sity34:118.2009
Specimen examined. PortugAl, Lisbon,Oeiras,EstaçãoAgronómicaNacional, stem of Foeniculum vulgare, 14Aug.2007,J.M. Santos(ex-typeculturesCBS123212=Di-C001/5, CBS123213=Di-C001/3).
Notes—Thisspecies(clade21)wasdescribedin2009onsenescent stems of Foeniculum vulgare(wildfennel)inPortugalbySantos&Phillips(2009).
Diaporthe manihotiaPunith.,Kavaka3:29.1976(1975) =Phomopsis manihotis Swarup,L.S.Chauhan&Tripathi,Mycopathol.Mycol.Appl.28,4:345.1966.
Specimen examined.rwAndA, on leaves of Manihot utilissima,9July1976, J. Semal(CBS505.76).
Notes — Phomopsis manihotis(clade25asD. manihotia)causesleafspotofcassava(Manihot esculenta),thoughthedis-easeisalsoreferredtoasPhomopsisblightoftapioca.Severeinfectionleadstodefoliationandstemlesions.Affectedareasbecome shrivelled with numerous pycnidia embedded in the tissue.Onseverelyinfectedstemsthebarkstartstograduallypeeloff,leadingtopartialortotalgirdling.ThediseaseisknownfromAfrica(Ethiopia,Nigeria),Asia(India),CentralAmericaandWestIndies(S.E.Dominica),andSouthAmerica(Colombia) (Sarbhoyetal. 1971,Mathur 1979,Farr&Rossman2012).
Diaporthe mayteni R.R.Gomes,C.Glienke&Crous,sp. nov. —MycoBankMB802939;Fig.11
Etymology.Namedafter thehost genus fromwhich itwas collected,Maytenus.
Conidiomata pycnidial, globose, immersed, scattered and ag-gregated,browntoblack,ostiolate,70–230μmwide,40–150μmtall,withshortnecks,40–140μm;outersurfacesmoothor covered in hyphae; pycnidal wall consisting of brown,thick-walled cells of textura angularis;conidialmassgloboseor exuding in cirrhi; predominantly yellow, pale luteous tocream.Conidiophores hyaline, subcylindrical to cylindrical, rarely branched above the septa, tapering towards the apex, 1–3-septate,(10–)13–27(–36)×(2–)3(–4)μm.Conidiogenous cells hyaline, subcylindrical, rarely tapering towards the apex, collarette present and not flared, with prominent periclinal thickening,(5–)6–10(–13)×2(–3)μm.Alpha conidia hyaline, oblongtoellipsoid,apexbluntlyrounded,baseobtuse;bigut-tulate,(5–)6(–7)×(2–)3μm.Beta and gamma conidiaabsent. Culturecharacteristics—ColoniesonPDAflat,withentireedge, cottony, olivaceous buff, with primrose aerial mycelium inconcentricrings,witholivaceouspatches;coloniesreach-ing66mmdiamafter2wkat25°C;reverseolivaceousbuffandgreenisholivaceous.OnOAflat,withentireedge,cottony
25R.R.Gomesetal.:Diaporthe
appressed, buff, white, the center of the colony pale olivaceous-grey,patchesisabellineandluteous;coloniesreaching56mmdiam;reversebuffandpaleolivaceousgrey.OnMEAflat,withentire edge, aerial mycelium cottony, white to pale olivaceous greyorolivaceousbuff;coloniesreaching37mmdiam;reversehazel,ochreous,withpatchesgreenishblackandolivaceousblack.
Specimen examined. BrAzil, Paraná,Colombo, endophytic speciesisolated from petiole of Maytenus ilicifolia(popularnameEspinheiraSanta), July2007,R.R. Gomes(holotypeCBSH-21096,ex-type culture CBS133185=LGMF938=CPC20314).
Notes — Diaporthe mayteni(clade30)growsendophyticallyin Maytenus ilicifoliainBrazil.
Diaporthe megalospora Ellis&Everh.,Proc.Acad.Nat.Sci.Philadelphia42:235.1890
Specimen examined. unknown, from Sambucus canadensis,Sept.1927,L.E. Wehmeyer (CBS143.27).
Notes — Diaporthe megalospora (clade15) is knownonSambucus canadensisfromNorthAmerica(Wehmeyer1933,Hanlin1963,Farr&Rossman2012).Freshcollectionsarere-quiredtodesignateanepitype,andfixthegeneticapplicationofthename.
Diaporthe melonis Beraha&M.J.O’Brien,Phytopathol.Z.94,3:205.1979
=Phomopsis cucurbitae McKeen,Canad.J.Bot.35:46.1957.
Specimens examined.indonesiA, Java, Muneng,Exp.Station,on Glycine soja, Sept.1987, H. Vermeulen(CBS435.87).–USA, Texas, RioGrandeValley,on Cucumis melo,1978,L. Beraha & M.J. O’Brien(ex-isotypecultureCBS507.78,specimenderivedfromcultureCBSH-891).
Notes—Clade7representsD. melonis(Beraha&O’Brien1979), and contains theex-isotype culture, andone isolatepreviouslyidentifiedasD. phaseolorumvar.sojae (thoughthetwoisolatesarenotidentical). Diaporthe melonisisfrequentlyreportedonsoybean(Santosetal.2011).Phomopsis cucurbitae(treatedhereassynonym)isreportedtohaveacosmopoli-tan distribution, and to cause black rot disease of greenhouse
cucumbers (McKeen1957,Punithalingam&Holliday1975,Ohsawa&Kobayashi1989).
Diaporthe musigena Crous&R.G.Shivas,Persoonia26: 119.2011
Specimen examined.AustrAliA,Queensland,BrisbaneBotanicalGarden,on leaves of Musasp.,14July2009,P.W. Crous & R.G. Shivas(ex-typecultureCBS129519=CPC17026).
Notes—Clade84representsD. musigena, isolated from Musa sp.inAustralia(Crousetal.2011).
Diaporthe neilliaePeck,Ann.Rep.N.Y.StateMus.Nat.Hist.39:52.1887(1886)
Specimen examined.unknown, on Spiraeasp.,Sep.1927,L.E. Wehmeyer(CBS144.27).
Notes — Diaporthe neilliae (clade 60)was originally de-scribed from Spiraeasp.fromNorthAmerica.Theoriginofthepresentisolate,however,remainsunclear(presumablyNorthAmerica).
Diaporthe neoarctii R.R.Gomes,C.Glienke&Crous,sp. nov. —MycoBankMB802940,Fig.12
Etymology.NamedafteritssuperficialresemblancetoDiaporthe arctii.
Conidiomata pycnidial,ampulliformtofinger-like,aggregated,darkbrowntoblack,immersed,ostiolate,300–450μmwide,200–670μm tall,with prominent necks 240–560μm long,outersurfacecoveredwithhyphae; pycnidal wall consisting of brown, thick-walled cells of textura angularis;conidialmassglobose, pale yellow.Conidiophores hyaline, ampulliform to subcylindrical, filiform, branched above the septa, taperingtowardstheapex,rarelyseptate,(12–)13–17(–18)×(2–)3μm.Conidiogenous cellshyaline,subcylindrical,filiform,straight,tapering towards the apex, collarette flared, periclinal thick-eningprominent, (10–)11–13(–14)× (1.5–)2(–3)μm.Alpha conidia hyaline, fusoid, apex acute, base obtusely rounded to subtruncate,bi-tomulti-guttulate,(9–)11–13(–14)×3(–4)μm.Beta and gamma conidianotobserved.
Fig. 11 Diaporthe mayteni(CBS133185).a.ConidiomatasporulatingonPNA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall; d,e.conidiogenouscells;f.betaconidia;g.alphaconidia.—Scalebars:b=85µm,allothers=10µm.
26 Persoonia–Volume31,2013
Culture characteristics — Colonies with sparse aerial my-celium,coveringthedishafter2wkinthedarkat25°C.OnMEAumberwithpatchesofgreyishsepia,umberinreverse.OnPDAfuscous-blackonsurfaceandinreverse.
Specimen examined. USA, NewJersey,isolatedfromAmbrosia trifida, May2001,G. Bills (holotypeCBSH-21094,ex-typecultureCBS109490 = GB6421=AR3450).
Notes—IsolatesoriginallyidentifiedasD. arctii cluster in clades19and67(Fig.1).Diaporthe neoarctii(clade16)wasisolated from Ambrosia trifidainNewJersey,USA,anddiffersmorphologically from the extype culture of D. arctii (alphaconidia7×3–3.5µm)(clade19).BasedonthesedifferencesD. neoarctii is described as a novel species.
Diaporthe nobilis complex Specimens examined.germAny,Münster,onstemofLaurus nobilis,Feb.1939,Kotthoff (CBS200.39). – JAPAn, isolate from Pinus pentaphyllabonzaiplantimportedfromJapanintotheNetherlands,May1979,G.H. Boerema (CBSH-16732,cultureCBS587.79).–koreA,onimportedchestnuts(Castanea sativa),collectedingrocerystoreinSydney,Australia,5July1999,K.A. Seifert(CBS113470=DAOM226800).–lAtviA, Rhododendron sp., I. Apine(CBS129167).–new zeAlAnd, on bark of Malus pumila, G.J. Samuels(CBS124030=GJS77-49);Waikatoregion,onPyrus pyrifolia, 2001, isol.W. Kandula,det.L. Castlebury(CBS116953=NZ-26,CBS116954=NZ-27).–yugoslAviA, on Hedera helix,July1989,M. MuntañolaCvetkovic (CBS338.89).
Notes—Clade62 is poorly resolved in this dataset, buthas some internal structure, suggesting that it contains several potentiallydistinctspecies.Moreisolateswouldberequiredtoresolve their taxonomy. Isolates in thiscladewereoriginallyidentifiedasPhomopsis fukushii (onPyrus pyrifolia,NewZea-land),P. conorum (onPinus pentaphylla, theNetherlands),P. castanea (onCastanea sativa,UK),Diaporthe perniciosa (Malus pumila,NewZealand),D. pulla(onHedera helix, Yu-goslavia)andD. nobilis(onLaurus nobilis,Germany).
Diaporthe nomurai Hara,inHara,Diseasesofcultivatedplants:140.1925.—Fig.13
Conidiomata in culture on OA sporulating poorly, globose, up to 300µmdiam,black,erumpent;creamconidialdropletsexudingfromcentralostioles;wallsconsistingof3–6layersofmediumbrown textura angularis.Conidiophores hyaline, smooth, 0–1- septate, rarely branched, densely aggregated, cylindrical, straight to sinuous, 10–20 × 2–3µm.Conidiogenous cells 6–10× 1.5–3µm,phialidic,cylindrical,terminal,withslighttaperto-wardsapex,1–1.5µmdiam,withvisiblepericlinalthickening;collarettenotflared,minute.Paraphysesnotobserved.Alpha conidia aseptate, hyaline, smooth, guttulate, fusoid-ellipsoid to clavate, straight to variously curved, tapering towards both ends,straight,apexsubobtuse,basetruncate,(7–)9–11(–13)× (2.5–)3µm.Gamma conidianotobserved.Beta conidia spindle-
Fig. 12 Diaporthe neoarctii (CBS109490).a.ConidiomatasporulatingonPDA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d–f.conidiogenouscells;g.alphaconidia.—Scalebars:b=225µm,allothers=10µm.
Fig. 13 Diaporthe nomurai (CBS157.29).a.ConidiomatasporulatingonPDA;b–e.conidiogenouscells;f.alphaconidia;g.betaconidia.—Scalebars=10µm.
27R.R.Gomesetal.:Diaporthe
shaped, aseptate, smooth, hyaline, apex acutely rounded, base truncate, tapering from lower third towards apex, gently curved, (20–)25–27(–30)×1.5(–2)µm. Culturecharacteristics—Coloniesreachingupto8cmdiamafter 2wk in thedarkat 25°C.OnMEAsurface isabelline,reversesepia.OnOAsurfacepalemousegreywithconcentricringsofmousegrey;reversemousegrey.OnPDAsurfaceandreversefuscous-black,withsparseaerialmycelium.
Specimen examined.JAPAn, on Morussp.,Dec.1929,K. Togashi(CBS157.29).
Notes—Clade58representsD. nomurai from Morus sp. in Japan.Diaporthe nomurai is known from hosts such as Morus alba, M. bombycis, M. latifolia and Morus sp.(Farr&Rossman2012).
Diaporthe novem J.M.Santos,Vrandečić&A.J.L.Phillips,Persoonia27:14.2011
=Phomopsis sp.9vanRensburgetal.,Stud.Mycol.55:65.2006.
Specimens examined. BrAzil, endophytic in petiole on Maytenus ilicifolia, July2007,R.R. Gomes(LGMF943=CPC20319).–CroAtiA, Slavonija,inseed on Glycine max, Sept.2008,T. Duvnjak(holotypeCBSH-20462,ex-typeculturesCBS127270=4-27/3-1,CBS127271=5/27/3-3,CBS127269=5-27/3-1).–romAniA,Calugareni,Distr.Mizil,livingleavesonPolygonatum odoratum,31July1970,O. Constantinescu(CBS354.71).
Notes — Clade 22 represents D. novem(Santosetal.2011),andcontainsanendophyticisolate(LGMF43)fromMaytenus ilicifolia,oneisolatepreviouslyidentifiedasDiaporthe pardalota on Polygonatum odoratumfromRomania,andthreeisolatesof D. novemwhichincludestheex-typeisolate.IsolateLGMF943representshighergeneticvariationthantheotherisolates,andappearstorepresentadifferentspecies.Sincethisisolatedid not sporulate, further morphological characterisation was not possible and we refrain from excluding it from the species pendingcollectionofmorestrainstoclarifyitsstatus.Diaporthe novem was reported as pathogen of Aspalathus linearis (vanRensburgetal.2006)asPhomopsis sp.9.Itwasrecently described as pathogen of Glycine max (Santosetal.2011).Thisspecieswas also reported on Hydrangea macrophylla (Santosetal.2010),Helianthus annuus and Vitis vinifera
(Santosetal.2011). ItisknowntooccurinBrazil,Romania,Croatia,Italy(Rekabetal.2004),Portugal(Santosetal.2010)andSouthAfrica(vanNiekerketal.2005,vanRensburgetal.2006).
Diaporthe oncostoma(Duby)Fuckel,Jahrb.NassauischenVereinsNaturk.23–24:205.1870.(1869–1870).—Fig.14
Basionym. Sphaeria oncostomaDuby,inRabenh.,Klotzsch.Herb.VivumMycol.:no.253.1854.
Conidiomata pycnidial, globose to ellipsoidal, aggregated as well as scattered, dark brown to black, immersed, ostiolate, 430–1170μmwide,370–790μmtall,lackingnecks,withoutersurfacecoveredinbrownhyphae;pycnidalwallconsistingofbrown, thick-walled cells of textura angularis; conidialmassgloboseorexudingincirrhi,whitetopaleluteousorpaleyellow.Conidiophores hyaline, subcylindrical, branched above the sep-ta,taperingtowardstheapex,1–2-septate,(10–)11–19(–22)× 3(–4)μm.Conidiogenous cells hyaline, subcylindrical, straight or curved, tapering towards the apex, collarette not flared, periclinal thickening prominent, (6–)7–9(–10)× (2–)3 μm.Alpha conidia hyaline, fusoid to ellipsoidal, straight to slightly curved, acute at apex, subobtuse at base, bi- or multi-guttulate, (7.5–)9–11(–12)× (2–)3(–4) μm.Gamma conidia hyaline, smooth, ellipsoid-fusoid, apex acutely rounded, and tapering towardstruncatebase,(11–)12–16×3(–3.5)µm.Beta conidia andsexualmorphnotobservedinculture. Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C.OnMEAsurfacedirtywhitewithprofuseaerialmycelium, reverse umber.OnOAsurface dirtywhitewithpatchesofumber,sameinreverse.OnPDAsurfaceandreversesienna,withsparseaerialmycelium.
Specimens examined.frAnCe,HteSavoie,Aigueblanche-Bellecombe,outletofriverMorelinIsère,ondeadbranchesofRobinia pseudoacacia,17July1978,H.A. van der Aa(CBS589.78).–germAny,Wolfenbüttel,onleafspot of Robinia pseudoacacia,15Nov.1996,H. Butin(CBS100454);Berlin,on leaf of Ilex aquifolium,Nov.1985,M. Hesse (CBS809.85).–russiA, on Robinia pseudoacacia,June2000,L. Vasilyeva(CBS109741=AR3445).
Notes — Diaporthe oncostoma(clade70)hasbeenconsid-ered to be a saprobic, or low virulence pathogen, which plays some role in natural pruning and self-thinning of black locust
Fig. 14 Diaporthe oncostoma (CBS100454).a.ConidiomatasporulatingonOA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d–f.conidiogenouscells;g.alphaconidia.—Scalebars:b=225µm,allothers=10µm.
28 Persoonia–Volume31,2013
forests (Robinia pseudoacacia) (Vajna2002).However, thisfungus has been reported as a causal agent of canker and severe dieback disease of black locust inRussia (Scerbin-Parfenenko1953)andinGreece(Michalopoulos-Skarmoutsos&Skarmoutsos1999).AlthoughisolateCBS809.85wasobtainedfrom Ilex aquifolium inGermany,we treat itasbelonging toD. oncostoma, as it matches the other strains phylogenetically as well as morphologically.
Diaporthe oxe R.R.Gomes,C.Glienke&Crous,sp. nov. — MycoBankMB802941;Fig.15
Etymology. Theword‘oxe’isanexpressionusedinnortheasternBrazilthatmeansamazementorsurprise,inrelationtothenumberofnovelspeciesisolatedasendophytesfrommedicinalplantsinBrazil.
Conidiomatapycnidialampulliformtofinger-like,eustromatic,convoluted to unilocular, semi-immersed, scattered, dark brown toblack,ostiolate,60–170μmwide,60–220μmtall;necksvariable in length, 20–150μm, outer surface coveredwithhyphae;pycnidalwallconsistingofbrown,thick-walledcellsoftextura angularis;conidialmassgloboseorexudingincirrhi,pale-luteoustocreamorpale-yellow.Conidiophores hyaline, ampulliform to subcylindrical, branched above the septa, taperingtowardstheapex,1–2-septate,(14–)17–25(–27)× (2–)3μm.Conidiogenous cellshyaline,subcylindrical,filiform,straight to curved, tapering towards the apex, collarette flared, periclinal thickeningprominent, (5–)6–10(–12)× 2(–3) μm.Alpha conidia hyaline, oblong to ellipsoid, apex bluntly rounded, baseobtusetosubtruncate,bi-tomulti-guttulate,(5–)6–7(–8)×(2–)3μm.Beta conidia hyaline, smooth, curved or hamate, (17–)22–30(–33)×2–3μm.Gamma conidianotobserved. Culture characteristics—Colonies onPDA flat,with anentire edge, surface mycelium dense and felty, ochreous to fulvous, dark brick, honey, buff, exudates rarely present as colourlessdrops;coloniesreaching49mmdiamafter2wkat25°C;reverseumber,ochreoustofulvous.OnOAflat,withanentire edge, surface mycelium dense and felty, rosy buff, pale olivaceous-grey, iron-grey, with patches olivaceous buff, exu-datesincolourlessandpaleluteousdrops;coloniesreaching40mmdiam;reversedarkbrick,olivaceous.OnMEAraised,with an entire edge, surface mycelium dense and felty, buff,
rosy-buff, with chestnut coloured exudates in the centre of the colony,andpaleluteousattheperiphery;coloniesreaching49mmdiam;reversechestnutandbay.
Specimens examined. BrAzil, on petiole of Maytenus ilicifolia,July2007,R.R. Gomes(holotypeCBSH-21098,ex-type cultureCBS133186=LGMF942=CPC20318);samecollectiondetails(CBS133187=LGMF936=CPC20312);onleafofSchinus terebinthifolius,July2007,J. Lima(LGMF915=CPC20291);onpetioleofM. ilicifolia, S.A.V. Pileggi(LGMF939=CPC20315);onpetioleofM. ilicifolia,July2007,R.R. Gomes(LGMF945=CPC20321).
Notes—Endophytic isolates (clade 34) frommedicinalplantsinBrazil.
Diaporthe padivar.padiG.H.Otth,Mitth.Naturf.Ges.Bern:99.1871(1870)
Specimens examined.sweden,Uppland,Dalbypar.,Tuna,onPrunus padus,17Apr.1988,K. & L. Holm(CBS114200=UPSC2569);Dalarna,Folkärnapar.,Sonnbo,onAlnus glutinosa,Dec.1992,K. & L. Holm(CBS114649=UPSC3496).
Notes — Diaporthe padi var. padi(clade56)representsaEuropean taxon occurring on Prunus.We chose the name D. padi over D. decorticans, as the basionym of the latter, Sphaeria decorticans,isanillegitimatehomonym.
Diaporthe paranensisR.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802942,Fig.16
Etymology.NamedafterParaná,thestateinBrazilfromwhereitwascollected.
Conidiomata pycnidial, ampulliform, semi-immersed, scattered, browntoblack,ostiolate,130–220μmwide,60–130μmtall;prominentnecks50–210μm long,outersurfacesmoothorcovered inhyphae;pycnidalwallconsistingofbrown, thick-walled cells of textura angularis;conidialmassglobose,pre- dominantly pale-luteous to yellow and some cases green-olivaceous.Conidiophores hyaline, subcylindrical to cylindri-cal,filiform,branchedabovetheseptaonaglobosecell,nottaperingtowardstheapex,2–3-septate,(14–)15–22(–26)× (2–)3(–4) μm.Conidiogenous cells hyaline, subcylindrical, filiform,rarelytaperingtowardstheapex,collarettepresentandflared,slightpericlinalthickening,(5–)8–14(–15)×2(–3)μm.
Fig. 15 Diaporthe oxe(CBS133186).a.ConidiomatasporulatingonPDA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d,e.conidiogenouscells;f.betaconidia;g.alphaconidia.—Scalebars:b=100µm,allothers=10µm.
29R.R.Gomesetal.:Diaporthe
Alpha conidia hyaline, fusoid-ellipsoidal, apex bluntly rounded, baseobtusetosubtruncate,bi-tomulti-guttulate,(6–)7–8(–9)×(2–)3μm.Beta conidia hyaline, smooth, curved or hamate and slightly curved, (16–)17–21(–23)× (1–)2 μm.Gamma conidianotobserved. Culture characteristics—Colonies onPDA flat,with anentire edge, mycelium growing in concentric rings, cottony texture,whitetosmoke-grey;coloniesreachingupto64mmdiamafter2wkat25°C;reversebuffandisabelline.OnOAflat, with an entire edge, aerial mycelium in concentric rings, ranging in colour from smoke-grey to grey-olivaceous and white in the centre; colonies reaching 44mmdiam; reverse iron-grey,grey-olivaceoustoolivaceous-buff.OnMEAflat,withanentire edge, aerial mycelium growing in concentric rings, with cottony texture, pale olivaceous-grey to grey-olivaceous and buff;coloniesreaching56mmdiam;reverseumber, fulvouswithpatchesofgreenishblack.
Specimen examined. BrAzil, Paraná, Colombo, endophytic species isolated from petiole of Maytenus ilicifolia(popularnameEspinheiraSanta),July2007,R.R. Gomes (holotypeCBSH-21099,ex-type culture CBS 133184=LGMF929=CPC20305).
Notes—Endophyticisolate(clade35)frommedicinalplantinBrazil.
Diaporthe perjunctaNiessl,Hedwigia15:153.1876 Specimen examined.AustriA, from Ulmus glabra,Oct.2001,A.Y. Rossman(ex-epitypecultureCBS109745=ARSEF3461=AR3461).
Notes — Diaporthe perjuncta(clade40)isassociatedwithfallen branches of Ulmus campestris and U. glabra (Ulmaceae).ThisspeciesisfoundinAustria,GermanyandPortu-gal.Diaporthe perjuncta is distinguished from D. viticola and D. australafricanabasedonmorphologyandDNAsequencedata(vanNiekerketal.2005).Pathogenicitystudiesanden-dophytic isolation of ‘D. perjuncta’fromgrapevinesinAustraliaand South Africa in fact represent isolates of D. australafricana (Mostertetal.2001a,Rawnsleyetal.2004,vanNiekerketal.2005).
Diaporthe perseae (Zerova)R.R.Gomes,C.Glienke&Crous,comb. nov.—MycoBankMB802944;Fig.17
Basionym.Phomopsis perseaeZerova,J.Bot.Acad.Sci.RSSUkraine1,1–2:307.1940.
Conidiomata pycnidialincultureonMEA,globose,upto400µmdiam,black,erumpent;creamconidialdropletsexudingfromcentralostioles;wallsconsistingof3–6layersofmediumbrowntextura angularis.Conidiophores hyaline, smooth, 1–3-septate, branched, densely aggregated, cylindrical, straight to sinuous, 15–35×3–4µm.Conidiogenous cells8–17×1.5–2.5µm,
Fig. 16 Diaporthe paranensis (CBS133184).a.ConidiomatasporulatingonPDA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d,e.conidiogenouscells;f.alphaandbetaconidia.—Scalebars:b=100µm,allothers=10µm.
Fig. 17 Diaporthe perseae (CBS151.73).a.ConidiomatasporulatingonPDA;b–d.conidiogenouscells;e.alphaandbetaconidia.—Scalebars=10µm.
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phialidic, cylindrical, terminal and lateral, with slight taper to-wardsapex,1–1.5µmdiam,withvisiblepericlinalthickening;col-laretteprominent,upto5µmlong.Paraphyses hyaline, smooth, subcylindricalwithobtuseends,2–4-septate,upto60µmlong, 3µmdiam.Alpha conidia aseptate, hyaline, smooth, guttulate, fusoid to ellipsoid, tapering towards both ends, straight, apex subobtuse, base subtruncate, (6–)7–8(–9)× 2(–2.5) µm.Gamma conidia aseptate, hyaline, smooth, ellipsoid-fusoid, apexacutelyrounded,basesubtruncate,9–14×1.5–2µm.Beta conidia spindle-shaped, aseptate, smooth, hyaline, apex acutely rounded, base truncate, tapering from lower third to-wardsapex,curved,(15–)22–25(–28)×1.5(–2)µm. Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C,withmoderateaerialmycelium.OnOAsurfaceochreous,withpatchesofdirtywhiteandiron-grey.OnPDAsurfacedirtywhitewithpatchesofsienna,reversesiennawithpatchesofumber.OnMEAsurfacesienna,withpatchesofumber,reverseumberwithpatchesofsienna.
Specimen examined.netherlAnds Antilles,Martinique,onyoungfruitofPersea gratissima,10July1972,E. Laville(CBS151.73).
Notes — Diaporthe perseae (clade86)wasoriginallyde-scribed from branches of dying Persea gratissimatreesinRus-sia.Basedonthemorphology(alphaconidia7–10.2×2.3–2.5µm;Uecker1988),thisstraincouldbeauthenticforthename.
Diaporthe phaseolorum (Cooke&Ellis)Sacc.,Syll.Fung.1: 692.1882
Basionym.Sphaeria phaseolorum Cooke & Ellis, Grevillea6, 39: 93.1878.
Specimens examined. BrAzil, endophytic in petiole on Maytenus ilicifolia,July2007, R.R. Gomes(LGMF927=CPC20303,LGMF941=CPC20317).–new zeAlAnd, from Olearia cf. rani, 22Jan.2003,G.J.M. Verkley (CBS113425);Actinidia chinensis, rotting fruit, kiwifruit orchard, S.R. Pennycook (CBS127465=GJS83-379).–unknown,Apr.1980,L. Beraha(CBS257.80).–USA,Mississippi, from Caperonia palustris, Oct.2003, A. Mengistu (CBS116019); Mississippi, from Aster exilis, Oct.2003,A. Mengistu(CBS116020).
Notes — Clade 4 represents isolates of D. phaseolorum.It includes two endophytic isolates fromMaytenus ilicifolia collected in Brazil, one isolate previouslymisidentified as D. melonis (CBS257.80),twoisolatesrespectivelyfromCaperonia palustres and Aster exilisintheUSA(Mengistuetal.2007),one isolate from Olearia cf.rami, and one from Actinidia chinensis.TheITSandTEF1sequencesofthiscladearesimilartosequences(GenBankU11323,U11373andEU222020,respec-tively)ofawell-characterisedisolateofD. phaseolorum(ATCC64802=FAU458).Byaccepting this cladeas authentic for
D. phaseolorum,wefollowtheprecedentsetbyvanRensburgetal.(2006),Mengistuetal.(2007)andSantosetal.(2011).
Diaporthe pseudomangiferae R.R.Gomes,C.Glienke&Crous, sp. nov.—MycoBankMB802945;Fig.18
Etymology.Namedafteritsmorphologicalsimilarityto Phomopsis mangiferae.
Conidiomatapycnidial,erumpent tosuperficialonPDA,glo-bose, up to 300 µm diam with elongated necks with central osti- olesthatexudeyellow-orangetocreamconidialdroplets;wallsof6–8layersofbrowntextura angularis.Conidiophores hyaline, smooth, 1–3-septate, branched, densely aggregated, cylindri-cal, straight to sinuous, 20–30 ×2–2.5µm.Conidiogenous cells phialidic, cylindrical, terminal and lateral with slight apical taper,10–15×2–3µm;collaretteflared,upto3µmlong.Paraphyses hyaline, smooth, cylindrical, septate, extending above conidiophores, straight to flexuous, unbranched or branched below,upto80µmlong,2–3µmwideatbase.Alpha conidia aseptate, hyaline, smooth, guttulate to granular, fusiform, taper-ing towards both ends, apex acutely rounded, base truncate, (6–)7–9(–10)×(2–)2.5(–3)µm.Beta and gamma conidia not seen(descriptionbasedonCBS101339). Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C,withmoderateaerialmycelium.OnOAsurfaceandreversedirtywhitewithpatchesofiron-grey.OnPDAsurfacedirtywhitetoochreous,reverseumber.OnMEAsurface greyish sepia with patches of iron-grey, reverse greyish sepiawithpatchesofiron-grey.
Specimens examined.dominiCAn rePuBliC, from Mangifera indica, P. de Leeuw,ATO-DLO,Wageningen (holotypeCBSH-21105, culture ex-typeCBS101339).–mexiCo, on fruit peel of Mangifera indica(CBS388.89).
Notes—Althoughtheseisolates(clade82)wereoriginallydescribed as representative of P. magiferae (dead leavesofMangifera indica,Pakistan),theydifferinhavinglargerconidio-mata,longerconidiophoresandlargeralphaconidia.
Diaporthe pseudophoenicicolaR.R.Gomes,C.Glienke& Crous, sp. nov.—MycoBankMB803839;Fig.19
Etymology.Namedafteritsmorphologicalsimilarityto Diaporthe phoenicicola.
ConidiomatapycnidialonMEA,upto400µmdiam,erumpent,globosewithneck;ostioleexudingyellow-orangeconidialdrop-lets;wallsconsistingof3–6layersofmediumbrowntextura angularis.Conidiophores hyaline, smooth, densely aggregated, 1–3-septate,branched,cylindrical,straighttocurved,12–45× 1.5–3µm.Conidiogenous cells phialidic, cylindrical, terminal
Fig. 18 Diaporthe pseudomangiferae (CBS101339).a.ConidiomatasporulatingonPNA;b.conidiomatasporulatingonPDA;c,d.conidiogenouscells; e.betaconidia;f.alphaconidia.—Scalebars=10µm.
31R.R.Gomesetal.:Diaporthe
and lateral with slight apical taper, 12–20 ×1.5–2µm,withvisible periclinal thickening; collarette flared, 2–5µm long.Paraphyses hyaline, smooth, cylindrical, 1–3-septate, extend-ing above conidiophores, straight to flexuous, unbranched or branched,upto100µmlong,and3µmwideatbase.Alpha conidia aseptate, hyaline, granular, smooth, fusiform, tapering towards both ends, straight, acutely rounded apex, and truncate base,(6–)7–8(–9)×(2–)2.5(–3)µm.Beta and gamma conidia notseen(descriptionbasedonCBS462.69). Culture characteristics — Colonies covering dish after 2 wk in thedark at 25°C,with sparseaerialmycelium.OnMEAsurface dirty white with patches of sienna, reverse umber with patchesofsienna.OnOAsurfacedirtywhitewithpatchesofsienna.OnPDAsurfaceochreouswithpatchesofolivaceous-grey,reverseiron-greywithpatchesofochreous.
Specimens examined.irAq,Prov.Basrah,ShaltElArab,showingdiebackon Mangifera indica,1976,M.S.A. AlMomen(CBS176.77).–sPAin,Mallorca,Can Pastilla, dead tops of green leaves on Phoenix dactylifera,27May1969,H.A. van der Aa(holotypeCBSH-21106,cultureex-typeCBS462.69).
Notes — Diaporthe pseudophoenicicola(clade89)isdistinctfrom D. phoenicicola(conidia8–12×2–2.5µm;Uecker1988)byhavingshorter,andwideralphaconidia.Asimilarstrainwasisolated from Mangifera indicainIraq(CBS176.77),suggestingthatthisspecieshasawiderhostrange.
Diaporthe pustulata Sacc.,Syll.Fung.(Abellini)1:610. 1882 Specimens examined.AustriA, on Acer pseudoplatanus,Oct.2001,A.Y. Rossman(CBS109742=AR3430andCBS109760=AR3535);Raab,AuWald,onPrunus padus,Oct.2001,A.Y. Rossman(CBS109784=AR3419).
Notes — Clade 44 contains one isolate from Prunus padus and two isolates from Acer pseudoplatanus, all isolated from Austria.Clade56containsanother isolateonPrunus padus fromSweden.ClearlytherearetwodifferentspeciesfromPrunus,oneisolatedinAustriaandanotherinSweden.BecauseD. pustulata was originally described on Acer pseudoplatanus, wetentativelyapplythisnametoisolatesinclade44.Toclarifythestatusofisolatesinclades44and56,however,additionalisolates and a comparison with type materials would be re-quired.
Diaporthe raonikayaporumR.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802947;Fig.20
Etymology. Raoni + Kayapo = afterthenameofaleader(Raoni)oftheindigenous KayapoethnictribeinBrazil.TheKayaposareinhabitantsoftheAmazonregioninBrazil.TheyusethemedicinalplantSpondias mombin, from which this species was isolated, as adornment or ornament, and for its medicinalproperties.
Conidiomata pycnidial, globose to conical or ampullifom, eus-tromatic and convoluted or unilocular, scattered, dark brown to black,immersed,ostiolate,110–200μmwide,50–130μmtall,
Fig. 19 Diaporthe pseudophoenicicola(CBS462.69).a,b.ConidiomatasporulatingonPDA;c,d.conidiogenouscells;e.alphaconidia.—Scalebars=10µm.
Fig. 20 Diaporthe raonikayaporum (CBS133182).a.ConidiomatasporulatingonPNA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d.conidiogenouscells;e.betawithafewalphaconidia;f.alphaconidia.—Scalebars:b=100µm,allothers=10µm.
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withprominentnecks40–140μmlong,outersurfacesmoothorcoveredinhyphae;pycnidalwallconsistingofbrown,thick-walled cells of textura angularis;conidialmassgloboseorexud-ingincirrhi,whitetopale-luteous.Conidiophores hyaline, am-pulliformtosubcylindrical,filiform,branchedabovethesepta,taperingtowardstheapex,1–3-septate,(16–)17–22(–26)× (2–)3μm.Conidiogenous cellshyaline,subcylindrical,filiform,straight to curved, tapering towards the apex, collarette not flared,periclinalthickeningprominent,(5–)7–9(–10)×(2–)3μm.Alpha and gamma conidia are formed in the same coni-diogenous cells.Alpha conidia hyaline, oblong to ellipsoid, apex bluntly rounded, base obtuse to subtruncate, bi- to multi-guttulate,(6–)7(–8)×(2–)3μm.Beta conidianotobserved.Gamma conidia hyaline, fusoid to subcylindrical, slightly curved, apex bluntly rounded, base obtuse to subtruncate, bi- to multi-guttulate,oreguttulate,(7–)9–11(–13)×(1–)2μm. Culture characteristics—Colonies onPDA flat,with anentire edge, aerial mycelium forming concentric rings with cot-tonytexture,olivaceous-buff,isabellinetohoneyonsurface;coloniesreaching63mmdiamafter2wkat25°C;reversepalepurplishgreytosmoke-grey.OnOAflat,withanentireedge,aerial mycelium forming concentric rings, white, olivaceous onsurface,coloniesreaching31mmdiam;reversebuffandgreenisholivaceous.OnMEAflat,withalobateedge,aerialmycelium forming wooly concentric rings, olivaceous-grey, greenish olivaceous and patches of amber on surface, colonies reaching51mmdiam;reversebrown-vinaceous.
Specimen examined. BrAzil,Pará,Redenção,endophyticspeciesisolatedfrom leaf of Spondias mombin(popularnameCajazeiraandTaperebá), July2007,K. Rodriguez(holotypeCBSH-21097,ex-type cultureCBS133182=LGMF923=CPC20299).
Notes—Endophyticisolate(clade31)frommedicinalplantinBrazil.
Diaporthe rhoina Feltgen,Vorstud.Pilzfl.Luxemb.,Nachtr.III: 145.1903
Specimen examined.unknown, on Rhus toxicodendron,Sept.1927,L.E. Wehmeyer(CBS146.27).
Notes—Thisspecies(clade95)wasoriginallydescribedon Rhus typhinafromLuxembourg.Europeanisolatesofthis
pathogenwillneed tobecollected toconfirm the identityofCBS146.27,whichispresumablyofNorthAmericanorigin.
Diaporthe saccarata(J.C.Kang,L.Mostert&Crous)Crous,comb. nov.—MB802948
Basionym.Phomopsis saccarataJ.C.Kang,L.Mostert&Crous,Sydowia53,2:230.2001.
Specimen examined.south AfriCA,WesternCapeProvince,JonkershoekMountains,Stellenbosch,oncankersofProtea repens,Mar.1999,S. Denman(ex-typecultureCBS116311=CPC3743).
Note — Diaporthe saccarata(clade71)isknowntocausea canker disease on shoots of Protea repens in South Africa (Mostertetal.2001b).
Diaporthe schini R.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802949;Fig.21
Etymology.Namedafter the host genus fromwhich itwas isolated,Schinus.
Conidiomata pycnidial, eustromatic, multilocular, immersed to erumpent, ostiolate, dark brown to black, scattered or ag-gregated,80–270μmwide,70–240μmtall,prominentnecks70–220μmlong,outersurfacecoveredwithhyphae;pycnidalwall consisting of brown, thick-walled cells of textura angularis;conidialmassglobose,pale-luteoustocream.Conidiophores hyaline,subcylindrical, filiform, rarelybranched, tapering to-wards theapex,0–1-septate, (11–)12–17(–20)× (2–)3(–4)μm.Conidiogenous cellshyaline,subcylindricalandfiliform,straight, tapering towards the apex, collarette not observed, withprominentpericlinalthickening5–6(–7)×(1–)2μm.Beta conidiahyaline,smooth,curvedorhamate(14–)22–28(–30)×(1–)2μm.Alpha and gamma conidianotobserved. Culturecharacteristics—ColoniesonPDAflat,witha lo-bate margin, surface mycelium sparse, felty and appressed, buff,honeytoisabelline;coloniesreaching30mmdiamafter2wk at 25°C; reverse greyish sepia, smoke-grey.OnOAwith a lobate margin, surface mycelium flat, sparse, felty and appressed,smoke-grey,olivaceous-grey,orolivaceousbuff;colonies reaching21mmdiam; reversepalemouse-grey to
Fig. 21 Diaporthe schini (CBS133181).a.ConidiomatasporulatingonPDA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d,e.conidiogenouscells;f.betaconidia.—Scalebars:b=135µm,allothers=10µm.
33R.R.Gomesetal.:Diaporthe
olivaceous-greyorbuff.OnMEAwithalobatemargin,surfacemycelium flat, dense, felty and appressed, buff with umber patches;coloniesreaching30mmdiam;reversedarkmouse-grey,umber,withpatchesofisabellineorluteous.
Specimen examined. BrAzil, Paraná,Curitiba, endophytic species isolated from leaf of Schinus terebinthifolius(popularnameAroeira),July2007,J. Lima (holotypeCBSH-21093,cultureex-typeCBS133181=LGMF921=CPC20297);samecollectiondetails(LGMF910).
Notes — Other than D. schini (clade11),additional endo-phytic isolates were also obtained from Schinus terebinthifolius inBrazil,butthesearemorphologicallydifferentandclusterinclades5and9(D. endophytica and D. terebinthifolii).
Diaporthe sclerotioides(Kesteren)Udayanga,Crous&K.D.Hyde,Fung.Diversity56:166.2012
Basionym.Phomopsis sclerotioidesKesteren,Neth.Jl.Pl.Path.73:115.1967.
Specimens examined.netherlAnds,Maarssen,onrootofCucumis sativus,June1967,H.A. van der Kesteren(ex-typecultureCBS296.67=ATCC18585=IMI151828=PD68/690);Roermond,onrootofC. sativus,Dec.1976(CBS710.76=PD76/674).
Notes — Diaporthe sclerotioides(clade28)wasoriginallyde-scribed from roots of Cucumis sativusintheNetherlands.Thisspecieshassubsequentlybeenreportedtocauseblackrootrot of Citrullus lanatus, Cucurmis sativus, C. ficifolia, C. maxi ma and C. moschata invariouscountriesintheworld(Uday-angaetal.2011).
Diaporthe scobinaNitschke,PyrenomycetesGermanici2: 293.1870
=Phomopsis scobina Höhn.,Sber.Akad.Wiss.Wien,Math.-naturw.Kl.,Abt.1115:681(33ofrepr.).1906.
Specimen examined.sCotlAnd, living and dead twig of Fraxinus excelsior, Feb.1938,J.A. MacDonald(CBS251.38).
Notes—Clade38 is representedbyD. scobina isolated from Fraxinus excelsiorinScotland.ThefungusisknownonthishostfromScotlandandPoland(Mulenkoetal.2008,Farr&Rossman2012).
Diaporthe sojaeLehman,Ann.MissouriBot.Gard.10:128.1923
≡ Diaporthe phaseolorumvar.sojae(Lehman)Wehm.,ThegenusDia-portheNitschkeanditssegregates47:1933. = Phomopsis longicollaHobbs,Mycologia77:542.1985. ≡ Diaporthe longicolla (Hobbs)J.M.Santos,Vrandečić&A.J.L.Phillips,Persoonia27:13.2011.
Specimens examined.CroAtiA, on Glycine max stem,Sept.2005,K. Vran dečić(specimenCBSH-20460,cultureCBS127267).–itAly, Bologna, from Glycine soja,1986, P. Giunchi(specimenCBSH-16776,cultureCBS100.87).– unknown, on Glycine soja (‘Blackhawk’)maturestem,A.A. Hildebrand(CBS180.55= ATCC12050=CECT2024). – USA,Mississippi,from Euphorbia nutans, A. Mengistu(CBS116017=DP0508)andfromGlycine max,Oct.2003, A. Mengistu(CBS116023);onGlycine soja seedling, J. Marcinkowska (CBS659.78=NRRL13656).
Notes—IsolatesofD. phaseolorum var.sojae clustered in twodistinctsclades(clade1,D. sojae;clade7,D. melonis).Diaporthe sojae causes pod and stem blight of soybean, while P. longicollaisknowntocauseseeddecay(Santosetal.2011).Severalauthorshavefounditdifficulttodistinguishthembasedon disease symptoms alone, and usually report them together (Almeida&Seixas2010).Hobbsetal.(1985)describedP. longicolla as a different species to D. sojae(Diaporthe phaseolorum var.sojae)basedonmorphologicalcharacters.Bothsymptomtypes, however, have also been linked to the same species
(Kulik1984,Morgan-Jones1989,Kulik&Sinclair1999).Con-sideringtheirgeneticsimilaritybasedonthefivegenesstudiedhere, disease etiology and common host, it appears that these isolates belong to the same species, which is distinct from D. phaseolorum(clade4).Diaporthe sojae(clade1)isanoldername than D. longicolla,andisthereforeappliedtothisclade.
Diaporthe sp. 1 Specimens examined. BrAzil,EMBRAPA/PR, on Glycine max seed, A. Almeida(LGMF947=CPC20323).–germAny, Bielefeld, human abscess, K. Plechulla (CBS119639= B11861).
Notes—Isolatesfromclade2appeartorepresentanovelspecies, Diaporthe sp. 1 (sterile). It is representedbyCBS119639,isolatedfromanabscessofamalepatientinGermany,andisolateLGMF947,obtainedfromsoybeanseedsinBrazil.Isolatesfromthiscladesharealowgenetichomologytoisolatesoftheclade4(D. phaseolorum;Fig.1,part1).Diaporthe species commonly described from soybean were alsoreportedasopportunistichumanpathogens.In1999,aspecies of Phomopsis was reported as etiological agent of a subcutaneousinfectiononthefingerofanimmunosuppressedfarmer and this genus was added to the list of fungi capable to causehumandisease(Suttonetal.1999).In2011,D. sojae (asPhomopsis longicolla),aknownpathogenofsoybean,wasidentifiedascausingskininfectioninanimmunocompromisedpatientafterkidneytransplantation.Theauthorsbelievedthatthispatientacquiredthefungusatleast5yrbefore,whenhehadcontactwithseedsorsoybeanplantsinEquatorialGuinea(Garcia-Reyneetal.2011).Another phytopathogenic species also described in soybean, Diaporthe phaseolorum, was reported causing osteomyelitis in patients with positive serology for human lymphotropic virus type1(HTLV-1),disturbingtheimmuneresponse.Thepatientwasafarmerandinoculationoccurredpossiblythroughinjurywith Amaranthus spinosusthorns(Iriartetal.2011).
Diaporthe sp. 2 Specimen examined.BrAzil, on petiole of Maytenus ilicifolia,July2007,R.R. Gomes(LGMF932=CPC20308).
Notes — Sterile, endophytic isolate from medicinal plant in Brazil,whichappearstorepresentanovelspecies(clade29).
Diaporthe sp. 3 Specimen examined.sCotlAnd, on Pseudotsuga menziesii,Mar.1929,G.G. Hahn(CBS287.29).
Notes — Clade 32 was tentatively named Diaporthe sp.3,andisrepresentedbyasingleisolatepreviouslyidentifiedasPhomopsis conorum, and obtained from Pseudotsuga menziesii inScotland.Thiscladewasnotresolved,becausethereareatleast eight different conifer species without any extype cultures (Udayangaetal.2011).
Diaporthe sp. 4 Specimen examined.BrAzil, endophytic in petiole on Maytenus ilicifolia, July2007,R.R. Gomes(LGMF944=CPC20320).
Notes—Sterileendophyticisolate(clade33)fromamedi-cinalplant inBrazil, appearing to representanundescribedspecies.
34 Persoonia–Volume31,2013
Diaporthe sp. 5 Specimen examined.itAly, from Acer opalus, W. Jaklitsch (CBS125575).
Notes—Thisisolate(clade37)representsanovelspeciesoccurring on Acer, which will be treated separately as part of anotherstudy(W.Jaklitsch,perscomm.).
Diaporthe sp. 6 Specimens examined.hong kong,UniversityDrive,onfruitofMaesa perlarius,18Dec.2000,K.D. Hyde(CBS115595=HKUCC10129,CBS115584=HKUCC7784).
Notes—The two strains (clade 85) studied herewereoriginallyidentifiedasP. pittospori(describedfromPittosporum twigs,USA,California),whichseemshighlyunlikely,astheywere isolated from fruit of Maesa perlarius inHongKong.Unfortunatelybothstrainsprovedtobesterile,sotheiridentitycouldnotbeconfirmed.
Diaporthe sp. 7 Specimen examined.indiA, Bangalore, on Anacardium occidentale,Aug.1978,H.C. Govindu (CBS458.78).
Notes—The identity of thepresent isolate (identifiedasPhomopsis anacardii)couldnotbeconfirmed,asthecultureproved tobesterile.However,phylogenetically (clade88) itrepresents a distinct taxon from D. anacardii(clade69),andwhenrecollected,shouldbedescribedasnew.
Diaporthe sp. 8 Specimen examined.BrAzil, from Aspidosperma tomentosum, K. Rodriguez(LGMF925=CPC20301).
Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C,withmoderateaerialmycelium.OnPDAsurfaceochreous,reversepale luteous.OnOAsurfaceandreverseluteous.OnMEAsurfacepaleluteous,reverseorangetoapricot.
Notes—Althoughthisisolate(clade90)appearstorepre-sent an undescribed species based on phylogenetic data, it provedtobesterile.Aswepresentlyonlyhaveasinglestrainofthistaxon,itstreatmentwillhavetoawaitfurthercollections.
Diaporthe stictica (Berk.&Broome)R.R.Gomes,C.Glienke& Crous, comb. nov.—MycoBankMB802950
Basionym.Phoma sticticaBerk.&Broome,Ann.Mag.Nat.Hist.,ser.II5:370.1850. ≡ Phomopsis stictica(Berk.&Broome)Traverso,Fl.Ital.Crypt.2,1:276.1906.
Specimen examined.itAly, Perugia, on dead twig of Buxus sempervirens, Dec.1954,M. Ribaldi(CBS370.54).
Notes — Diaporthe stictica(clade74)representsaEuropeanspecies occurring on Buxus sempervirens (Italy,Germany).Although the present isolate could be authentic for the name, thiscouldnotbeconfirmedbasedonmorphology,astheisolateprovedtobesterile.
Diaporthe subordinaria(Desm.)R.R.Gomes,C.Glienke&Crous, comb. nov.—MycoBankMB802951
Basionym.Phoma subordinariaDesm.,Ann.Sci.Nat.,Bot.ser.3,9:284.1849. ≡ Phomopsis subordinaria(Desm.)Traverso,Fl.Ital.Crypt.Pars1:Fungi.Pyrenomycetae.Xylariaceae,Valsaceae,Ceratostomataceae:232.1906.
Specimens examined.new zeAlAnd, blackened seed of Plantago lanceolata,Apr.1999,B. Alexander(CBS101711).–south AfriCA, Eastern Cape Province,Grahamstown, on stalks ofPlantago lanceolata, 2Dec. 1989, R. Shivas (CBS464.90).
Notes — Diaporthe subordinaria (clade 18) has a globaldistribution on Plantago lanceolata, on which it causes a stalk disease(deNooij&vanderAa1987).Itispossiblethatthedis-ease relates to several different species occurring on Plantago, but this matter can only be resolved following futher collections andcorrelationwithtypematerial.
Diaporthe tecomaeSacc.&P.Syd.,Syll.Fung.14:550.1899(nom.nov.forD. interrupta Niessl).—Fig.22
?= Phoma tecomae Sacc.,NuovoGiorn.Bot.Ital.8:201.1876. ≡ Phomopsis tecomae (Sacc.)Traverso&Spessa,Bol.Soc.Brot.Coim-bra,sér.1,25:124.1910.
Conidiomata pycnidial, sporulating poorly on OA, globose, up to1mmdiam,black,erumpent,multilocular;creamconidialdropletsexudingfromcentralostioles;wallsconsistingof3–6layers of medium brown textura angularis.Conidiophores hya-line in upper region, pale brown at base, smooth, 1–3-septate, branched, densely aggregated, cylindrical, straight to sinuous, 20–30 ×2–3μm.Conidiogenous cells8–15×1.5–3μm,phia-lidic, cylindrical, terminal and lateral, with slight taper towards apex,1μmdiam,withvisiblepericlinalthickening;collarettenotflared,minute.Paraphysesnotobserved.Beta conidia spindle-shaped, aseptate, smooth, hyaline, apex acutely rounded, base truncate, tapering from lower third towards apex, apex strongly curved, (17–)22–24(–26)×1.5(–2)μm.Alpha and gamma conidianotobserved. Culture characteristics — Colonies covering dish after 2 wk inthedarkat25°C.OnOAfluffy,dirtywhitewithpatchesofgreyolivaceous.OnPDAdirtywhitewithpatchesofolivaceousgrey and isabelline, reverse with patches of dirty white, brown vinaceousanddarkbrick.OnMEAdirtywhitewithpatchesofisabelline and olivaceous grey, reverse brown vinaceous with patchesofdarkbrick.
Fig. 22 Diaporthe tecomae(CBS100547).a.ConidiomataformingonPNA;b,c.conidiogenouscells;d.betaconidia.—Scalebars=10µm.
35R.R.Gomesetal.:Diaporthe
Specimen examined.BrAzil,SaoPaulo,SerradaMantiqueira, mycoceci-dium caused by Prosopodium tecomicola on living young branch of Tabebuia sp.,27Sept.1997,coll.A. Aptroot,isol.H.A. van der Aa(specimenCBSH-16834,cultureCBS100547).
Notes — Diaporthe tecomae was a new name proposed for D. interruptaNiessl(onTecoma radicans,Portugal),astheepi-thetwasalreadyoccupied.ThelinkbetweentheDiaporthe and Phomopsisstateremainstobeproven.Theasexualmorphwasoriginally described as Phoma tecomae (fromItalyonTecoma radicans, conidiophores 20 ×1µm,conidia8×3µm;Saccardo1878),andisprobablydistinctfromthefungusrepresentedbyCBS100547,whichoccursonTabebuia sp.inBrazil.However,as no ex-type strains are available of D. tecomae (clade10),and no alpha conidia were observed in culture, this could not beconfirmed,andispendingfreshcollections.
Diaporthe terebinthifolii R.R.Gomes,C.Glienke&Crous,sp. nov.—MycoBankMB802952;Fig.23
Etymology. Named after the host species from which it was isolated, Schinus terebinthifolius.
Conidiomata pycnidial, globose to conical, immersed, ostiolate, brown to black, scatteredor aggregated, 95–110μmwide,140–160µmtall,rarelyformsnecks,butwhenpresent,theyareshortandcoveredwithhyphae;pycnidalwallconsistingof brown, thick-walled cells of textura angularis;conidial mass globose,whiteorpale-luteoustocream.Conidiophores hyaline, subcylindrical,filiform,branchedabovesepta,taperingtowardstheapex, 1–2-septate, (13–)15–21(–22)× 2(–3)μm.Beta conidiogenous cells hyaline, ampulliform to subcylindrical and filiform,taperingtowardstheapex,collarettepresentandnotflared,slightpericlinalthickening,(3–)6–10(–14)×2(–3)μm.Beta conidiahyaline,smooth,curvedorhamate,(18–)20–24 (–26)×1(–2)μm.Alfa and gamma conidianotobserved. Culturecharacteristics—ColoniesonPDAflat,withanentireedge, aerial mycelium cottony, greyish white, colonies reaching 64mmdiamafter2wkinthedarkat25°C;reversebuff.OnOAflat, entire edge, aerial mycelium cottony, with concentric rings, pale olivaceous-grey, smoke-grey and greyish white, colonies
reaching48mmdiam;reverseolivaceous-greyandolivaceousbuff.OnMEAflat,withanentireedge;aerialmyceliumcottony,smoke-grey, colonies reaching60mmdiam; reverseumberwithpatchesoffuscous-black.
Specimens examined.BrAzil,Paraná,Curitiba,endophyticspeciesiso-lated from leaf of Schinus terebinthifolius(popularnameAroeira),July2007,J. Lima (holotypeCBSH-21097,ex-typecultureCBS133180 =LGMF914=CPC20290);samecollectiondetails(LGMF909=CPC20285,LGMF907=CPC20283,LGMF913=CPC20289).
Notes—Themultigene analysis of isolates in clade 9exhibitedinsignificanthomologytosequencesfoundinGen-Bank.AnisolatepreviouslyidentifiedasPhomopsis tecomae (CBS100547)alsoresidesinthisclade,butismorphologicallydistinct. No morphologically similar isolates are known from S. terebinthifolius, and thus we designate these isolates as representativeofanewtaxon.
Diaporthe toxica P.M.Will.,Highet,W.Gams&Sivasith.,Mycol.Res.98:1367.1994
Specimens examined.western AustrAliA,Morawa,onstemofLupinus angustifolius,6May1991,J.B. Nunn(ex-typecultureCBS534.93=ATCC96741);Serpentine,onLupinussp.,8June1993,P.M. Williamson (CBS535.93);Medina,onLupinussp.,8June1993,P.M. Williamson(CBS546.93).
Notes—Clade45containsthreeisolatesofD. toxica, in-cludingtheex-typeculture(CBS534.93),isolatedfromLupinus angustifoliusinWesternAustralia.TwovarietiesofPhomopsis (P. leptostromiformis var. leptostromiformis and P. leptostromiformisvar.occidentalis)wereidentifiedascausingdiseasein Lupinussp.Diaporthe woodii was later recognised as the sexual state of P. leptostromiformisvar.occientalis(Punithal-ingam1974),whileWilliamsonet al. (1994) designated thename D. toxica for the sexual state of the toxicogenic variety, P. leptostromiformisvar.leptostromiformis.Lupins (Lupinusspp.)aregrown inmanypartsof theworldasagrainlegumecrop.Theseedsareusedforanimalfeedandincreasinglyasflourforhumanconsumption.Theplantsincreasesoilnitrogenandaregrowninrotationwithothercrops.
Fig. 23 Diaporthe terebinthifolii(CBS133180).a.ConidiomatasporulatingonPNA;b,c.transversesectionthroughconidiomata,showingconidiomatalwall;d,e.conidiogenouscells;f.betaconidia.—Scalebars:b=100µm,c=25µm,allothers=10µm.
36 Persoonia–Volume31,2013
InAustraliathestubbleleftafterharvestingaidssoilconserva-tionand isavaluablesummer feed for livestock.Diaporthe toxica is considered to be an important limiting factor to more extensivesowingoflupins.Thisorganismhasbeenreportedtocausestemblightinyounglupins(Lupinus luteus)(Ostazeski&Wells1960)andtoproducephomopsins(Culvenoretal.1977).Thesemycotoxinscause theanimal liverdiseaseknownaslupinosis(Gardiner1975,Allen&Wood1979).
Diaporthe vacciniiShear,U.S.Dept.Agric.Tech.Bull.258:7. 1931
=Phomopsis vacciniiShear,U.S.Dept.Agric.Tech.Bull.258:7.1931.
Specimens examined.USA,Massachusetts,onOxycoccus macrocarpos, Mar.1932,C.L. Shear(ex-typecultureCBS160.32=IFO32646);Michigan,on Vaccinium corymbosum, G.C. Adams (CBS118571);NewJersey,on V. macrocarpon,1988,L. Carris (CBS122112=FAU474);Michigan,on V. corymbosum,1992,D.C. Ramsdell(CBS122114=FAU634,CBS122115=FAU590);NorthCarolina,fromV. corymbosum,pre-1999,D.F. Farr(CBS122116=DF5022).
Notes—Clade63 consists of six isolatesofD. vaccinii, includingtheex-typestrain(CBS160.32)isolatedonVaccinium corymbosumfromtheUSA.Diaporthe vaccinii causes fruit rot and twig blight and leaf spots of Vacciniumspp.(blueberries)intheUSA(Alfierietal.1984,Farretal.2002,Farr&Rossman2012).TheprincipalhostsareAmericanandEuropeancran-berries(Vaccinium macrocarpon, V. oxycoccos, V. oxycoccos var. intermedium),highbushblueberry(V. corymbosum)andrabbiteyeblueberry(V. ashei).Diaporthe vaccinii is restricted to cultivated Vacciniumspecies.ThewildEuropeanspecies,V. oxycoccos, which usually occurs in mountain bogs, could beapotentialreservoirforthepest.IntheEPPOregionithasbeenreportedfromRomania(foundinexperimentalplotsofintroducedAmericancultivars,butdidnotestablish(Teodorescuetal.1985)),UK(foundinplantsoriginallyimportedfromtheNetherlandsandUSA,butdidnotestablish(Wilcox&Falconer1961,Baker1972)).Symptoms in susceptible blueberry cultivars include blighting of1-yr-oldwoodystemswithflowerbuds.Infectedsucculent,current-year shoots wilt in 4 d and become covered with minute lesions.Thefunguscontinuestotraveldownwardthroughthestem,killingmajorbranches,andoftenentireplants(Wilcox1939,Daykin&Milholland1990).Infectedfruitsturnreddish-brown,soft,mushy,oftensplittingandcausingleakageofjuice(Milholland&Daykin1983).
Diaporthe vexans(Sacc.&P.Syd.)Gratz,Phytopathology32: 542.1942
Basionym.Phoma vexansSacc.&P.Syd.,Syll.Fung.(Abellini)14,2:889.1899. ≡ Phomopsis vexans(Sacc.&P.Syd.)Harter,J.Agric.Res.2,5:338.1914.
Specimen examined.USA,from Solanum melongena,Dec.1914,L.L. Harter(CBS127.14).
Notes — Diaporthe vexans(clade12)causesfruitrot,leafspot,stemandtipblightdiseaseofeggplants(Solanum melongena and S. wendlandii)andothersolanaceousspecies,Acacia sp.(Fabaceae),Prunussp.(Rosaceae)andSorghum bicolor (Poaceae),Capsicum annuum and Lycopersicon esculentum (Solanaceae).Thedisease iswidespread inNorthAmerica,theWestIndies,andEasternandCentralAsia,alsoinAfrica(Senegal,Tanzania,Zambia)andMauritius(Punithalingam&Holliday1972).AdditionalrecordsincludeBrunei,Haiti,Iran,IraqandRomania(Harter1914,Farr&Rossman2012).
Diaporthe viticola Nitschke, PyrenomycetesGermanici 2:264.1870
Specimens examined.AustriA, Vienna,Risenbergbach-Weg, onLaburnum anagyroides,May2001,A.R. Rossman(CBS109492).–CAnAdA, British Columbia, Sidney, on Epilobium angustifolium,Oct.2001,M. Barr (CBS109768=AR3478).–frAnCe,Dordogne,nearSarlatlaCanéda,1-yr-old stems on Asphodelus albus,20May1995,G. Verkley(CBS759.95).– netherlAnds,Utrecht,Baarn,inbranchesandtwigsofAucuba japonica, Jan.1995,G. Verkley(CBS106.95);onRosa rugosa,18Mar.1985,G.H. Boerema(CBS266.85=PD85/25);Lelystad,indeadstemon Lupinussp.,May1982,H.A. van der Aa(CBS449.82);Wieringermeer,Robbenoordbos,in dead stem on Lupinus arboreus,12Mar.1991,H.A. van der Aa & F. Meurs (CBS312.91);Flevoland,treesinfrontofInfoCentreLepelaarsplassen,inleaf spot on Fraxinus excelsior,31Aug.1997,H.A. van der Aa(CBS100170);Baarn, in dead stem on Dipsacus fullonum,14June1985,H.A. van der Aa (CBS502.85);ontwigonSalixsp.,Apr.1962,G.H. Boerema(CBS446.62).– PortugAl, on Vitis vinifera(Galegodurado),1Jan.1998,A.J.L. Phillips (CBS114011=CPC2677);Burgaes,SantoTirso,onVitis vinífera,16Feb.1998,A.J.L. Phillips(ex-type cultureCBS113201=CPC5683).–sweden, Skåne,Maglehempar.,onSambucuscf.racemosa, 14Apr.1989,K. Holm & L. Holm(CBS114436=UPSC2960).–UK,Sheffield,onA. japonica,July1996,G. Verkley(CBS794.96).
Notes — Diaporthe viticola(clade50)isknownfromseveralhosts, but especially from grapevines, on which it causes a canespotdiseaseinEurope(Portugal,Germany).Merrinetal.(1995)referredtoseveralAustralianisolatesfromgrapevinesas Phomopsis taxon1.The samespecieswas reportedbyPhillips(1999)asD. perjunctaandbyScheperetal.(2000)asD. viticola.Inasubsequentstudy,Mostertetal.(2001a)chosetofollowPhillips(1999)andappliedthenameD. perjuncta to taxon 1. However, theyalsonoted thatminormorphologicaldifferences existed in perithecia and ascospores between the EuropeanandSouthernHemispherematerial,which led tothe description of a novel taxon, D. australafricana, for isolates fromAustraliaandSouthAfrica(vanNiekerketal.2005),andtheepitypificationofD. viticolabasedonEuropeanmaterial.Based on the results obtained here, D. viticola (clade50) isclosely related to D. australafricana(clade49),andisclearlydistinguishable from D. perjuncta(clade40).
Diaporthe woodiiPunith.,Mycol.Pap.136:51.1974 =Phomopsis leptostromiformisvar.occidentalis,R.G.Shivas,J.G.Allen&P.M.Will.,Mycol.Res.95:322.1991.
Specimen examined.western AustrAliA,Medina,stemsofLupinussp.,8July1993,P.M. Williamson(CBSH-5319,cultureCBS558.93).
Notes—Clade94representsDiaporthe woodii(CBS558.93),whichwascharacterisedbyWilliamsonetal.(1994),basedontheex-typestrain(IMI166508).Diaporthe crotalariae (clade92), D. aspalathi (clade93) and D. woodii arecloselyrelatedspecies.Diaporthe woodii causes stem rot, stem cankers, leaf infections and seed decay of Lupinus angustifolius and L. cosenfinii, and blight and seed discoloration of L. albus, L. angustifolius, L. cosentinii, L. luteus, L. pilosus and Trifolium subterraneum (subterraneanclover).ThefungusisknowntooccurinBrazil,SouthAfrica,USA(Florida),andWesternAustralia(Williamsonetal.1994).
Diaporthe woolworthii(Peck)Sacc.,Syll.Fung.(Abellini)1:615. 1882
Basionym.Valsa woolworthiiPeck,Ann.Rep.N.Y.StateMus.Nat.Hist.28:73.1876(1875).
Specimen examined.unknown, on Ulmus americana,Sept.1927,L.E. Wehmeyer(CBS148.27).
Notes—Clade57containsasingleisolateofD. woolworthii from Ulmus americana. This taxon represents anAmericanspecies occurring on Ulmus,sothisculture(presumablyfromNorthAmerica),couldprovetobeauthenticforthename.
37R.R.Gomesetal.:Diaporthe
DISCUSSION
Amajoraimofthepresentstudywastoresolvethetaxonomyof Diaporthe species occurring on diverse hosts, either as pathogens, saprobes, or as harmless endophytes.To de-limitate these taxa, nine genes were screened, from which the bestfivewereselectedtoconductamulti-genephylogeneticanalysis(ITS,TEF1,ACT,HISandCAL).Diaporthe represents a highly complex genus containing numerous cryptic species, several of which are newly described in the present study, while others remain unclear, awaiting fresh collections and type studies.ManyDiaporthe species that are morphologically similar proved to be genetically distinct, and several isolates thatwereformerlyidentifiedbasedontheirhost,wereshowntorepresentdifferenttaxa.Although the genera Diaporthe and Phomopsis have received much taxonomic attention, few phylogenetic studies have thus far been conducted, and hence the taxonomy of this group is stillproblematic.Duetothelackofreferencestrains,andthefactthatfewgenelociotherthanITShaveinthepastbeenusedforDNAanalysis,mostoftheconclusionsreachedthusfar have been incorrect, meaning that published literature will havetobeinterpretedwithcare.Inthisstudywestudied15endophyticDiaporthe species from Brazil.Threewere not identified to species level, twowereidentifiedasD. novem and D. phaseolorum, while a further 10weredescribedasnew.Highgeneticdiversitywasfoundamongsttheanalysedisolatesfrommedicinalplants.Speciesof Diaporthe are commonly isolated as endophytes from sev-eralhostsintemperateandtropicalregions(Bussabanetal.2001,Muralietal.2006,Rossmanetal.2007,Botella&Diez2011,González&Tello2011).Skaltsasetal.(2011)isolated108Diaporthe isolates from asymptomatic leaves and bark of threedifferent hosts (Hevea brasiliensis, H. guianensis and Micandra spp.) fromCameroon,Mexico andPeru.Usingamultigene approach, the authors found more than 40 phylo-genetic species, of which several appeared to represent novel taxa(Skaltsasetal.2011).DespitemembersofDiaporthe commonly being described as phytopathogenic, an increasing number of reports link this genus to endophytic studies, focusing on its potential as a producerofenzymesandnovelsecondarymetabolites,withantibiotic, fungicideandanticanceractivity (Dai et al. 2005,Elsaesseretal.2005,Linetal.2005,Silvaetal.2005,Wuetal.2008,Kumaran&Hur2009,Weber2009,Vesterlundetal.2011).TheecologyofspeciesofDiaporthe remains poorly understood, as some endophytes isolated from the medicinal plant Maytenus ilicifoliawereidentifiedasD. phaseolorum(clade4)andD. novem (clade22),respectivelyknowaspathogenofsoybean(Santosetal.2011)andAspalathus linearis (vanRensburgetal.2006).Diaporthe novem is also reported from hosts such as Hydrangea macrophylla (Santosetal.2010),Helianthus annuus and Vitis vinifera (Santosetal.2011).Thesereportsagreewiththepogostickhypothesis,postulatingthathost-specificfungalplantpathogensfrequentlyexhibittheabilitytocolonisenon-hosttis-sue,enablingthemtodispersefurther,inanattempttofindthehostonwhichtheyarepathogenic(Crous&Groenewald2005).ThetaxonomyofDiaporthe(incl.Phomopsis)hastraditionallybeen based on host association, with species being described ontheassumptionthattheyarehost-specific.Inthepresentstudy the taxonomy of all Diaporthe isolates deposited in the CBS culture collection over time were reviewed, based on this assumption.The employment of this criterion, has ledto an exponential growth in the number of taxa described in Diaporthe thus far (Uecker 1988).However, in spite of the
apparent synonymies outlined in this study, there was evidence for a huge proliferation of cryptic taxa that were formerly over-looked based on a morphological approach in the absence of molecular data.Speciesdelimitation inDiaporthe based on morphological characters is challenging, as most taxa in culture donotproduceallsporestatesoftheasexual(alpha,betaandgammaconidia)orthesexualmorph.Thedescriptionofnoveltaxa in Diaportheintheabsenceofmoleculardata(atleastITSandHISorTUB;seediscussioninnextsection)shouldthusbestronglydiscouragedinthefuture.Inconclusionthus,itseemsthatinspiteofthefactthatthesetaxareadilycoloniseorco-colonisenon-hosts(seealsoRehner&Uecker1994,Mostertetal.2001a,Farretal.2002,Diogoetal.2010),thereisstillamultitudeofundescribedtaxaawaitingfurtherstudyinthiscomplex.Itisthushopedthatthephylo-genetic backbone generated here provides a stable platform to enable future studies by others interested in the biology of Diaporthe.
Phylogenetic species recognition by genealogical concordance Tayloretal.(2000)developedtheGenealogicalConcordancePhylogeneticSpeciesRecognition(GCPSR)concepttodefinethe limits of sexual species, using the phylogenetic concor-danceofmultipleunlinkedgenes.This concept hasprovedgreatlyusefulinfungi,becauseitismorefinelydiscriminatingthan other species concepts, as several species are unable to be crossed, or cannot be recognised due to the lack of distin-guishingmorphologicalcharactersorsterility(Reynolds1993,Tayloretal.2000,Caietal.2011).Theadoptionofgenealogicalconcordance for species recognition in Diaporthe enabled us to distinguish species that were otherwise not possible to identify duetoeithersterility,orthelossofspecificcharacterstates.For instance, D. viticola and D. australafricana are two closely relatedspecies (clades50and49, respectively)associatedwithgrapevines.Theyaremorphologicallysimilar,butoccurondifferentcontinents(vanNiekerketal.2005).Thesespecieshave probably accumulated genetic differences due to their geographicalisolation.SeveralcrypticspecieswererecentlydescribedinothergenerausingtheGCPSRcriterion,someof which are consistent with allopatric divergence, because these species occupy non-overlapping areas separated by geographicbarriers,e.g.inCladosporium(Benschetal.2012),Colletotrichum(Dammetal.2012a,b),Harknessia(Crousetal.2012),Ilyonectria(Cabraletal.2012a,b)andPhyllosticta (Glienkeetal.2011), tonamebuta few.UsingtheGCPSRconceptitispossibletodefinethegeneticvariationobservedinsomespecies,butstillinsufficienttoestablishthemasdis-tinctspecies,sincegeneticflowstilloccursbetweenthem.Forexample,isolatesofclades79–90clustereddifferentlybasedon analyses of the different genes, probably because of recent geneflowamongthem.Wehavecomparedthelocationandmonophylyofthestrainsineach clade in the phylogenetic tree of the combined alignment (Fig.1)tothosephylogenetictreesobtainedfromtheindividualloci to determine the species boundaries and species resolu-tion.ThefivelociselectedfortheBayesianphylogenyhaveasimilarresolutionforspeciesdiscrimination,rangingfromTEF1resolving72outofthe95species,toHISandTUBresolving84ofthe95species.TheITSregion,whichisoftenconsideredtobelessthanopti-mal for closely related species, was not much better or worse (resolving75ofthe95species)thantheotherincludedloci.However,given the recentacceptanceof the ITS regionasofficialfungalbarcode(Schochetal.2012)anditsintermedi-ate resolving power in the present study, this locus should not
38 Persoonia–Volume31,2013
bediscardedfromfuturestudies.Also,TEF1,whichhasinthepast been used as additional locus for phylogenetic studies of Diaporthe,performedtheworstinthisstudy(resolving72ofthe95species),althoughthiswasnotmuchworsethanITSandCAL(resolving75and74ofthe95species,respectively).TheHISandTUBregionsappeartohavethebestresolutionforspe-cies discrimination in the present study and therefore are good candidatesassecondarymarkerstothecommonlyusedITSregion.SimilarresultswerealsoreportedforITS,CAL,TEF1andTUBbyUdayangaetal.(2012),whosuggestedthatTUBbe considered as secondary phylogenetic marker for Diaporthe.
The importance of epitypification in Diaporthe Thebestoptiontosupplementpoortypematerial isviaepi-typification(Cannonetal.2012).ToemploytheGCPSRconceptinfungi,DNAismostlyextractedfrompoorlypreserved,ancientherbariumspecimenswithdifficulty,andinmanycasesitonlyresultsinshortsequencesoftheITSregion(Quaedvliegetal.2011,Cheewangkoonetal.2012).Therefore,epitypificationof living material, and its preservation and deposit in publically available collections and databases, are important steps to provide a stable platform to enable others to test future hy-potheses.Althoughitisnotaprerequisite,itisstronglyrecom-mended that the chosen epitype should originate from the same geographical region and host, and have morphological, cultural and pathological characteristics similar to those described in theoriginalpublication(seeDammetal.2012a,b,Cannonetal.2012,Weiretal.2012).Despitethefactthatcloseto2000speciesofDiaporthe(incl.Phomopsis)havebeendescribedinliterature,hardlyanyex-type strains areavailable today, themajority ofwhichwereincludedinthepresentstudy.Duetothelackofex-typestrains,the taxonomy of several species continue to be unresolved, someofwhichareimportantplantpathogens.Aseriouseffortwill thus be called for to recollect and redescribe all these old names.Analternativeapproachwouldbetosimplystartover,ensuring that all newly described names are based not only on morphology,butalsosupplementedbyDNAbarcodes.How-ever,aslongasfungalnomenclatureisgovernedbytheICN,thisseemsunobtainable.Eventually though,allmycologistswill realise that a stable fungal nomenclature must incorporate DNAdata,andthatthisisonlyachievableifmycologyfollowsacodeofnomenclaturethatincorporatesthisrequirement.
AcknowledgementsWethankthetechnicalstaff,ArienvanIperen(cul-tures),MarjanVermaas(photographicplates)andMiekeStarink-Willemse(DNAisolation,amplificationandsequencing)fortheirinvaluableassistance.Weare grateful to theBrazilian agencyCAPES for financial support toRenataR.Gomes.
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