Research Article |
Corresponding author: Xinlei Fan ( xinleifan@bjfu.edu.cn ) Academic editor: George Mugambi
© 2018 Xinlei Fan, Zhuo Du, Jadson D.P. Bezerra, Chengming Tian.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Fan X, Du Z, Bezerra JDP, Tian C (2018) Taxonomic circumscription of melanconis-like fungi causing canker disease in China. MycoKeys 42: 89-124. https://doi.org/10.3897/mycokeys.42.29634
|
Melanconis-like species comprise latent fungal pathogens with a wide range of woody hosts. Taxonomy of these pathogens is difficult due to their uninformative descriptions and similar asexual morphology. Based on molecular phylogenies, many species of this group were placed in various families of Diaporthales. In this study, eight species of melanconis-like fungi were isolated from Betula albosinensis, B. platyphylla (Betulaceae), Cornus controversa (Cornaceae), Corylus mandshurica (Betulaceae) and Juglans regia (Juglandaceae) in China. These species were phylogenetically placed in three families of Diaporhthales, i.e. Juglanconis juglandina, J. oblonga (Juglanconidaceae), Melanconiella betulicola sp. nov., M. corylina sp. nov. (Melanconiellaceae), Melanconis betulae, Ms. itoana, Ms. stilbostoma (Melanconidaceae) and one new genus, Sheathospora (Melanconiellaceae). Sheathospora is proposed to accommodate Melanconiella cornuta with conical and discrete pycnidia with aseptate, hyaline, cylindrical to ellipsoidal conidia with distinct hyaline sheath on branches of Cornus controversa. Combined analyses of ITS, LSU, CAL, RPB2 and TEF1-α sequence data were used to construct the molecular phylogeny. Additionally, we provided separate phylogenetic trees for three families (Juglanconidaceae, Melanconidaceae and Melanconiellaceae) to show the species distribution of melanconis-like fungi in China.
Diaporthales , phylogeny, taxonomy, wood-inhabiting fungi
Melanconium (Diaporthales) was introduced by
Molecular phylogenetics have had a major impact in taxonomic rearrangements of fungi since the early 1990s (
During trips to collect forest pathogens that cause canker or dieback diseases in China, several melanconis-like taxa associated with various disease symptoms were collected in Beijing, Gansu, Heilongjian, Jilin, Ningxia, Shaanxi and Tibet Provinces. As the higher-level phylogeny of many genera within the melanconis-like taxa remains largely unresolved in China, this project was initiated to address this issue. In this paper, we identified eight melanconis-like species residing in three families of Diaporthales; recognised three genera within Melanoconiellaceae; and described two new species in Melanconiella as well as one new genus to accommodate Melanconiella cornuta.
Fresh specimens of melanconis-like fungi were collected from infected branches of seven hosts during collection trips in China (Table
Species | Culture/strain/specimen | Location | Host | GenBank accession numbers | ||||
ITS | LSU | CAL | RPB2 | TEF1-α | ||||
Apiosporopsis carpinea | CBS 771.79 | Switzerland | Carpinus betulus | NA | AF277130 | NA | NA | NA |
Apiosporopsis sp. | 11Af2-1 | Japan | Alnus firma | NA | AB669034 | NA | NA | NA |
Apoharknessia insueta | CBS 111377 | Brazil | Eucalyptus pellita | JQ706083 | AY720814 | NA | NA | NA |
CBS 114575 | Colombia | Eucalyptus sp. | NA | AY720813 | NA | NA | NA | |
Asterosporium asterospermum | MFLU 15-3555 | Italy | Fagus sylvatica | NA | MF190062 | NA | MF377615 | NA |
CBS 112404 | Italy | Fagus sylvatica | NA | AB553745 | NA | NA | NA | |
KT2138 | Japan | Fagus crenata | NA | AB553744 | NA | NA | NA | |
Auratiopycnidiella tristaniopsidis | CBS 132180 = CPC 16371 | Australia | Tristaniopsis laurina | JQ685516 | JQ685522 | NA | NA | NA |
Cainiella johansonii | Kruys 731 | Sweden | Dryas octopetala | NA | JF701920 | NA | NA | NA |
Chapeckia nigrospora | AR 3809 | USA | Betula sp. | JF681957 | EU683068 | NA | NA | NA |
Chiangraiomyces bauhiniae | MFLUCC 17-1669 | Thailand | Bauhinia sp. | MF190118 | MF190064 | NA | MF377604 | NA |
MFLUCC 17-1670 | Thailand | Bauhinia sp. | MF190119 | MF190065 | NA | MF377603 | NA | |
Chrysocrypta corymbiae | CBS 132528 | Australia | Corymbia sp. | JX069867 | JX069851 | NA | NA | NA |
Coniella diplodiella | CBS 111858 = CPC 3708 | France | Vitis vinifera | AY339323 | AY339284 | NA | KX833423 | KX833603 |
Coniella koreana | CBS 143.97 | Korea | NA | KX833584 | AF408378 | NA | KX833490 | KX833684 |
Coniella musaiensis var. hibisci | AR 3534 = CBS 109757 | South Africa | Hibiscus sp. | KX833589 | AF408337 | NA | NA | KX833689 |
Coniella straminea | CBS 149.22 = CPC 3932 | USA | Fragaria sp. | AY339348 | AF362569 | NA | KX833506 | KX833704 |
Coniella wangiensis | CBS 132530 = CPC 19397 | Australia | Eucalyptus sp. | JX069873 | JX069857 | NA | KX833509 | KX833705 |
Coryneum depressum | AR 3897 | Austria | Quercus cerris | NA | EU683074 | NA | NA | NA |
Coryneum modonium | AR 3558 | Austria | Castanea sativa | NA | EU683073 | NA | NA | NA |
Coryneum umbonatum | AR 3541 | Austria | Quercus cerris | NA | EU683072 | NA | NA | NA |
MFLUCC 15-1110 | Italy | Quercus sp. | MF190121 | MF190067 | NA | MF377610 | NA | |
MFLUCC 13-0658 | Italy | Quercus sp. | MF190120 | MF190066 | NA | MF377609 | NA | |
Cryphonectria macrospora | AR 3444 = CBS 109764 | Russia | Quercus mongolica | EU199182 | AF408340 | NA | EU220029 | NA |
Cryphonectria nitschkei | AR 3433 = CBS109776 | Russia | Quercus mongolica | DQ120761 | AF408341 | NA | NA | NA |
Cryphonectria parasitica | ATCC 38755 | USA | Castanea dentata | AY141856 | EU199123 | NA | DQ862017 | EU222014 |
Cryptodiaporthe aesculi | AFTOL-ID 1238 = CBS 109765 | Austria | Aesculus hippocastanum | DQ323530 | AF408342 | NA | EU199138 | GU354004 |
AR3640 = CBS 121905 | USA | Aesculus hippocastanum | EU254994 | EU255164 | NA | EU219269 | DQ313558 | |
LCM 447.01 | Germany | Aesculus hippocastanum | GU367076 | NA | NA | GU367110 | GU354002 | |
Cryptosporella betulae | AR 3524 = CBS 109763 | Austria | Betula pendula | EU199180 | AF408375 | NA | EU199139 | EU221884 |
Cryptosporella hypodermia | AR 3552 | Austria | Ulmus minor | EU199181 | AF408346 | NA | EU199140 | NA |
Cryptosporella suffusa | AR 3496 = CBS 109750 | Austria | Alnus incana | EU199207 | AF408376 | NA | EU199163 | EU221945 |
Cytospora cenisia | AR 3522 = CBS 109752 | Austria | Juniperus communis | NA | AF408385 | NA | NA | NA |
Cytospora chrysosperma | CFCC 89600 | China | Sophora japonica | KR045623 | KR045623 | NA | KU710951 | KU710915 |
Cytospora elaeagni | CFCC 89633 | China | Elaeagnus angustifolia | KF765677 | KF765693 | NA | KU710956 | KU710919 |
Cytospora leucostoma | CFCC 50468 | China | Betula platyphylla | KT732949 | KT732968 | NA | NA | NA |
Cytospora nivea | AR 3512 | Austria | Salix purpurea | NA | AF408367 | NA | NA | NA |
Cytospora sacculus | AR 3416 = CBS 109756 | Russia | Quercus mongolica | NA | AF408386 | NA | NA | NA |
AR 3426 = CBS 109777 | Austria | Quercus robur | NA | AF408387 | NA | NA | NA | |
Dendrostoma mali | CFCC 52102 | China | Malus spectabilis | MG682072 | MG682012 | NA | MG682032 | MG682052 |
Dendrostoma osmanthi | CFCC 52106 | China | Osmanthus fragrans | MG682073 | MG682013 | NA | MG682033 | MG682053 |
CFCC 52107 | China | Osmanthus fragrans | MG682074 | MG682014 | NA | MG682034 | MG682054 | |
CFCC 52108 | China | Osmanthus fragrans | MG682075 | MG682015 | NA | MG682035 | MG682055 | |
CFCC 52109 | China | Osmanthus fragrans | MG682076 | MG682016 | NA | MG682036 | MG682056 | |
Dendrostoma quercinum | CFCC 52103 | China | Quercus acutissima | MG682077 | MG682017 | NA | MG682037 | MG682057 |
CFCC 52104 | China | Quercus acutissima | MG682078 | MG682018 | NA | MG682038 | MG682058 | |
CFCC 52105 | China | Quercus acutissima | MG682079 | MG682019 | NA | MG682039 | MG682059 | |
Diaporthe decedens | AR 3459 = CBS 109772 | Austria | Corylus avellana | KC343059 | AF408348 | NA | NA | NA |
Diaporthe detrusa | AR 3424 = CBS 109770 | Austria | Berberis vulgaris | KC343061 | AF408349 | NA | NA | KC343787 |
Diaporthe eres | AR 3538 = CBS 109767 | Austria | Acer campestre | KC343075 | AF408350 | NA | NA | KC343801 |
Diaporthella corylina | CBS 121124 | China | Corylus sp. | KC343004 | NA | NA | NA | NA |
Diaporthella sp. | CN5 | Italy | Corylus avellana | KP205483 | NA | NA | NA | NA |
CN13 | Italy | Corylus avellana | KP205484 | NA | NA | NA | NA | |
Diaporthosporella cercidicola | CFCC 51994 | China | Cercis chinensis | KY852492 | KY852515 | NA | NA | NA |
CFCC 51995 | China | Cercis chinensis | KY852493 | KY852516 | NA | NA | NA | |
CFCC 51996 | China | Cercis chinensis | KY852494 | KY852517 | NA | NA | NA | |
Diaporthostoma machili | CFCC 52100 | China | Machilus leptophylla | MG682080 | MG682020 | NA | MG682040 | MG682060 |
CFCC 52101 | China | Machilus leptophylla | MG682081 | MG682021 | NA | MG682041 | MG682061 | |
Disculoides eucalypti | CPC 17650 | Australia | Eucalyptus sp. | JQ685517 | JQ685523 | NA | NA | NA |
Disculoides eucalyptorum | CBS 132184 = CPC 17648 | Australia | Eucalyptus viminalis | NR120090 | JQ685524 | NA | NA | NA |
Ditopella ditopa | AR 3423 = CBS 109748 | Austria | Alnus glutinosa | EU199187 | EU199126 | NA | EU199145 | NA |
Erythrogloeum hymenaeae | CPC 18819 | Brazil | Hymenaea courbaril | JQ685519 | JQ685525 | NA | NA | NA |
Gnomonia gnomon | CBS 199.53 | Italy | Corylus avellana | AY818956 | AF408361 | NA | EU219295 | EU221885 |
Harknessia eucalypti | CBS 342.97 | Australia | Eucalyptus regnans | AY720745 | AF408363 | NA | NA | NA |
Harknessia leucospermi | CBS 775.97 | South Africa | Leucospermum sp. | NR137147 | AY720824 | NA | NA | NA |
Harknessia molokaiensis | AR 3578 = CBS 109779 | USA | Eucalyptus robusta | NA | AF408390 | NA | NA | NA |
Harknessia syzygii | CBS 111124 = CPC184 | South Africa | Syzygium cordatum | AY720738 | AY720834 | NA | NA | NA |
Hercospora tiliae | AR 3526 | Austria | Tilia tomentosa | NA | AF408365 | NA | NA | NA |
Hyaliappendispora galii | MFLUCC 16-1208 | Italy | Galium sp. | MF190149 | MF190095 | NA | NA | NA |
Involutscutellula rubra | CBS 192.71 | Japan | Quercus phillyraeoides | MG591899 | MG591993 | NA | MG976476 | MG592086 |
Juglanconis appendiculata | D140 | Greece | Juglans nigra | KY427138 | KY427138 | NA | KY427188 | KY427207 |
D96 | Austria | Juglans nigra | KY427139 | KY427139 | NA | KY427189 | KY427208 | |
D96A | Austria | Juglans nigra | KY427140 | KY427140 | NA | KY427190 | KY427209 | |
MC | Greece | Juglans nigra | KY427141 | KY427141 | KY427242 | KY427191 | KY427210 | |
MC2 | Spain | Juglans nigra | KY427142 | KY427142 | KY427243 | KY427192 | KY427211 | |
MC4 | Spain | Juglans nigra | KY427143 | KY427143 | KY427244 | KY427193 | KY427212 | |
ME17 | Austria | Juglans nigra | KY427144 | KY427144 | KY427245 | KY427194 | KY427213 | |
Juglanconis juglandina | D142 | Austria | Juglans nigra | KY427145 | KY427145 | NA | KY427195 | KY427214 |
CFCC 51727* | China | Juglans nigra | KY363854 | KY363859 | MK096394 | MK096439 | NA | |
CFCC 51728* | China | Juglans nigra | KY363855 | KY363860 | MK096395 | MK096440 | NA | |
CFCC 51729* | China | Juglans nigra | KY363856 | KY363861 | MK096396 | MK096441 | NA | |
MC1 | Austria | Juglans nigra | KY427146 | NA | KY427246 | KY427196 | KY427215 | |
MC3 | Spain | Juglans nigra | KY427147 | KY427146 | KY427247 | KY427197 | KY427216 | |
ME16 | Austria | Juglans nigra | KY427148 | KY427147 | KY427248 | KY427198 | KY427217 | |
ME22 | Austria | Juglans nigra | KY427149 | KY427148 | KY427249 | KY427199 | KY427218 | |
ME23 | Austria | Juglans nigra | KY427150 | KY427150 | KY427250 | KY427200 | KY427219 | |
Juglanconis oblonga | CFCC 51725* | China | Juglans nigra | KY363852 | KY363857 | MK096392 | MK096437 | NA |
CFCC 51726* | China | Juglans nigra | KY363853 | KY363858 | MK096393 | MK096438 | NA | |
ME14 | USA | Juglans cinerea | KY427151 | KY427151 | KY427251 | KY427201 | KY427220 | |
ME15 | USA | Juglans cinerea | KY427152 | KY427152 | KY427252 | KY427202 | KY427221 | |
ME18 | Japan | Juglans ailanthifolia | KY427153 | KY427153 | KY427253 | KY427203 | KY427222 | |
ME19 | Japan | Juglans ailanthifolia | KY427154 | KY427154 | KY427254 | KY427204 | KY427223 | |
Juglanconis pterocaryae | ME20 | Japan | Pterocarya rhoifolia | KY427155 | KY427155 | KY427255 | KY427205 | KY427224 |
Lamproconium desmazieri | MFLUCC 14-1047 | Russia | Tilia cordata | KX430132 | KX430133 | NA | NA | MF377592 |
MFLUCC 15-0870 | Russia | Tilia tomentosa | KX430134 | KX430135 | NA | MF377605 | MF377591 | |
Lasmenia sp. | CBS 124123 | Puerto Rico | Nephelium lappaceum | GU797406 | JF838338 | NA | NA | NA |
CBS 124124 | Puerto Rico | Nephelium lappaceum | JF838336 | JF838341 | NA | NA | NA | |
Luteocirrhus shearii | CBS 130776 | Australia | Banksia baxteri | NR120254 | NG042770 | NA | NA | NA |
Macrohilum eucalypti | CPC 10945 | New Zealand | Eucalyptus sp. | DQ195781 | DQ195793 | NA | NA | NA |
CPC 19421 | Australia | Eucalyptus piperita | KR873244 | KR873275 | NA | NA | NA | |
Melanconiella betulicola | CFCC 52482* | China | Betula albosinensis | MK096312 | MK096352 | NA | MK096397 | MK096272 |
CFCC 52483* | China | Betula albosinensis | MK096313 | MK096353 | NA | MK096398 | MK096273 | |
Melanconiella carpinicola | MNM | Austria | Carpinus betulus | JQ926232 | JQ926232 | NA | JQ926304 | JQ926370 |
MNUK | UK | Carpinus betulus | JQ926234 | JQ926234 | NA | JQ926306 | JQ926372 | |
MSMI | Austria | Carpinus betulus | JQ926235 | JQ926235 | NA | JQ926307 | JQ926373 | |
Melanconiella chrysodiscosporina | MCH | Austria | Carpinus betulus | JQ926238 | JQ926238 | NA | JQ926310 | JQ926376 |
MEE | Austria | Carpinus betulus | JQ926240 | JQ926240 | NA | JQ926312 | JQ926378 | |
MGG | Austria | Carpinus betulus | JQ926242 | JQ926242 | NA | JQ926314 | JQ926380 | |
Melanconiella chrysomelanconium | MCM | Austria | Carpinus betulus | JQ926247 | JQ926247 | NA | JQ926319 | JQ926385 |
MEUK | UK | Carpinus betulus | JQ926249 | JQ926249 | NA | JQ926321 | JQ926387 | |
MGUK | UK | Carpinus betulus | JQ926255 | JQ926255 | NA | JQ926327 | JQ926393 | |
Melanconiella chrysorientalis | MGB | Croatia | Carpinus orientalis | JQ926256 | JQ926256 | NA | JQ926328 | JQ926394 |
MGP | Croatia | Carpinus orientalis | JQ926257 | JQ926257 | NA | JQ926329 | JQ926395 | |
MVH | Croatia | Carpinus orientalis | JQ926259 | JQ926259 | NA | JQ926331 | JQ926397 | |
Melanconiella corylina | CFCC 52484* | China | Corylus mandshurica | MK096314 | MK096354 | NA | MK096399 | MK096274 |
CFCC 52485* | China | Corylus mandshurica | MK096315 | MK096355 | NA | MK096400 | MK096275 | |
Melanconiella decorahensis | CBS 159.26 | USA | Betula sp. | JQ926260 | JQ926260 | NA | JQ926332 | JQ926398 |
MD | France | Betula pendula | JQ926261 | JQ926261 | NA | JQ926333 | JQ926399 | |
MED | France | Betula pendula | JQ926262 | JQ926262 | NA | JQ926334 | JQ926400 | |
Melanconiella echinata | DAOM 121196 | USA | Carpinus caroliniana | JQ926263 | JQ926263 | NA | N/A | N/A |
Melanconiella elegans | AR 3830 | USA | Carpinus caroliniana | JQ926264 | JQ926264 | NA | JQ926335 | JQ926401 |
BPI 843574 | USA | Carpinus caroliniana | JQ926266 | JQ926266 | NA | JQ926337 | JQ926403 | |
BPI 872067 | USA | Carpinus caroliniana | JQ926267 | JQ926267 | NA | JQ926338 | JQ926404 | |
Melanconiella ellisii | BPI 843491 | USA | Carpinus caroliniana | JQ926268 | JQ926268 | NA | N/A | JQ926405 |
BPI 878343 | USA | Carpinus caroliniana | JQ926271 | JQ926271 | NA | JQ926339 | JQ926406 | |
BPI 883227 | USA | Carpinus caroliniana | JQ926269 | JQ926269 | NA | N/A | N/A | |
Melanconiella flavovirens | MFV1 | Austria | Corylus avellana | JQ926274 | JQ926274 | NA | JQ926342 | JQ926409 |
MFV2 | Austria | Corylus avellana | JQ926275 | JQ926275 | NA | JQ926343 | JQ926410 | |
MFV3 | Italy | Corylus avellana | JQ926276 | JQ926276 | NA | JQ926344 | JQ926411 | |
Melanconiella hyperopta | MCHBV | Austria | Carpinus betulus | JQ926280 | JQ926280 | NA | JQ926346 | JQ926413 |
MCR | Austria | Carpinus betulus | JQ926283 | JQ926283 | NA | JQ926349 | JQ926416 | |
MHG | Switzerland | Carpinus betulus | JQ926285 | JQ926285 | NA | JQ926351 | JQ926418 | |
Melanconiella hyperopta var. orientalis | MHP | Croatia | Carpinus orientalis | JQ926288 | JQ926288 | NA | JQ926352 | JQ926420 |
MHVA | Croatia | Carpinus orientalis | JQ926287 | JQ926287 | NA | JQ926353 | JQ926419 | |
MSK | Croatia | Carpinus orientalis | JQ926286 | JQ926286 | NA | JQ926354 | JQ926421 | |
Melanconiella meridionalis | MOA | Austria | Ostrya carpinifolia | JQ926289 | JQ926289 | NA | JQ926355 | JQ926422 |
MOK | Croatia | Ostrya carpinifolia | JQ926290 | JQ926290 | NA | JQ926356 | JQ926423 | |
MOM | Austria | Ostrya carpinifolia | JQ926291 | JQ926291 | NA | JQ926357 | JQ926424 | |
Melanconiella ostryae | CBS 208.38 | USA | Ostrya virginiana | JQ926297 | JQ926297 | NA | JQ926363 | JQ926430 |
Melanconiella spodiaea | MVS | Croatia | Carpinus orientalis | JQ926299 | JQ926299 | NA | JQ926365 | JQ926432 |
MSH | Austria | Carpinus betulus | JQ926298 | JQ926298 | NA | JQ926364 | JQ926431 | |
SPOD | Croatia | Carpinus betulus | JQ926300 | JQ926300 | NA | JQ926366 | JQ926433 | |
Melanconis alni | AR 3529 | Russia | Duschekia maximowiczii | NA | AF362566 | NA | NA | NA |
AR 3748 | Austria | Alnus viridis | EU199195 | EU199130 | NA | EU199153 | NA | |
AR 4016 = CBS 121480 | Austria | Alnus alnobetula | EU254863 | NA | NA | EU219298 | EU221894 | |
CBS 109773 | Austria | Alnus viridis | DQ323523 | AF408371 | NA | EU219300 | EU221896 | |
Melanconis betulae | CFCC 50471* | China | Betula albosinensis | KT732952 | KT732971 | NA | KT732984 | KT733001 |
CFCC 50472* | China | Betula albosinensis | KT732953 | KT732972 | NA | KT732985 | KT733002 | |
CFCC 50473* | China | Betula albosinensis | KT732954 | KT732973 | NA | KT732986 | KT733003 | |
Melanconis italica | MFLUCC 16-1199 | Italy | Alnus cordata | MF190151 | MF190096 | NA | NA | NA |
MFLUCC 17-1659 | Italy | Alnus cordata | MF190151 | MF190097 | NA | MF377602 | NA | |
Melanconis itoana | CFCC 50474* | China | Betula albosinensis | KT732955 | KT732974 | NA | KT732987 | KT733004 |
CFCC 52876* | China | Betula albosinensis | MK096324 | MK096364 | NA | MK096409 | MK096284 | |
CFCC 52877* | China | Betula albosinensis | MK096326 | MK096366 | NA | MK096411 | MK096286 | |
CFCC 52878* | China | Betula albosinensis | MK096327 | MK096367 | NA | MK096412 | MK096287 | |
MAFF 410080 | Japan | Betula ermanii | JX522738 | NA | NA | NA | NA | |
Melanconis marginalis | AR 3442 = CBS 109744 | Canada | Alnus rubra | EU199197 | AF408373 | NA | EU219301 | EU221991 |
MAFF 410218 | Japan | Alnus maximowiczii | JX522742 | NA | NA | NA | NA | |
Melanconis stilbostoma | CBS 109778 = AR 3501 | Austria | Betula pendula | DQ323524 | AF408374 | NA | EU219299 | EU221886 |
CBS 121894 = MS | NA | Betula pendula | JQ926229 | JQ926229 | NA | JQ926302 | JQ926368 | |
CFCC 50475* | China | Betula platyphylla | KT732956 | KT732975 | NA | KT732988 | KT733005 | |
CFCC 50476* | China | Betula platyphylla | KT732957 | KT732976 | NA | KT732989 | KT733006 | |
CFCC 50477* | China | Betula platyphylla | KT732958 | KT732977 | NA | KT732990 | KT733007 | |
CFCC 50478* | China | Betula platyphylla | KT732959 | KT732978 | NA | KT732991 | KT733008 | |
CFCC 50479* | China | Betula platyphylla | KT732960 | KT732979 | NA | KT732992 | KT733009 | |
CFCC 50480* | China | Betula platyphylla | KT732961 | KT732980 | NA | KT732993 | KT733010 | |
Melanconis stilbostoma | CFCC 50481* | China | Betula platyphylla | KT732962 | KT732981 | NA | KT732994 | KT733011 |
CFCC 50482* | China | Betula platyphylla | KT732963 | KT732982 | NA | KT732995 | KT733012 | |
CFCC 52843* | China | Betula platyphylla | MK096338 | MK096378 | NA | MK096423 | MK096298 | |
CFCC 52844* | China | Betula platyphylla | MK096341 | MK096381 | NA | MK096426 | MK096301 | |
CFCC 52845* | China | Betula platyphylla | MK096343 | MK096383 | NA | MK096428 | MK096303 | |
CFCC 52846* | China | Betula platyphylla | MK096347 | MK096387 | NA | MK096432 | MK096307 | |
CFCC 52847* | China | Betula platyphylla | MK096348 | MK096388 | NA | MK096433 | MK096308 | |
CFCC 52848* | China | Betula platyphylla | MK096349 | MK096389 | NA | MK096434 | MK096309 | |
CFCC 52849* | China | Betula platyphylla | MK096328 | MK096368 | NA | MK096413 | MK096288 | |
CFCC 52850* | China | Betula platyphylla | MK096329 | MK096369 | NA | MK096414 | MK096289 | |
CFCC 52851* | China | Betula platyphylla | MK096330 | MK096370 | NA | MK096415 | MK096290 | |
CFCC 52852* | China | Betula platyphylla | MK096331 | MK096371 | NA | MK096416 | MK096291 | |
CFCC 52853* | China | Betula platyphylla | MK096332 | MK096372 | NA | MK096417 | MK096292 | |
CFCC 52854* | China | Betula platyphylla | MK096333 | MK096373 | NA | MK096418 | MK096293 | |
CFCC 52855* | China | Betula platyphylla | MK096334 | MK096374 | NA | MK096419 | MK096294 | |
CFCC 52856* | China | Betula platyphylla | MK096335 | MK096375 | NA | MK096420 | MK096295 | |
CFCC 52857* | China | Betula platyphylla | MK096336 | MK096376 | NA | MK096421 | MK096296 | |
CFCC 52858* | China | Betula platyphylla | MK096337 | MK096377 | NA | MK096422 | MK096297 | |
CFCC 52859* | China | Betula platyphylla | MK096339 | MK096379 | NA | MK096424 | MK096299 | |
CFCC 52860* | China | Betula platyphylla | MK096340 | MK096380 | NA | MK096425 | MK096300 | |
CFCC 52861* | China | Betula platyphylla | MK096342 | MK096382 | NA | MK096427 | MK096302 | |
CFCC 52862* | China | Betula platyphylla | MK096344 | MK096384 | NA | MK096429 | MK096304 | |
CFCC 52863* | China | Betula platyphylla | MK096345 | MK096385 | NA | MK096430 | MK096305 | |
CFCC 52864* | China | Betula platyphylla | MK096346 | MK096386 | NA | MK096431 | MK096306 | |
CFCC 52865* | China | Betula platyphylla | MK096316 | MK096356 | NA | MK096401 | MK096276 | |
CFCC 52866* | China | Betula platyphylla | MK096317 | MK096357 | NA | MK096402 | MK096277 | |
CFCC 52867* | China | Betula platyphylla | MK096318 | MK096358 | NA | MK096403 | MK096278 | |
CFCC 52868* | China | Betula platyphylla | MK096319 | MK096359 | NA | MK096404 | MK096279 | |
CFCC 52869* | China | Betula platyphylla | MK096320 | MK096360 | NA | MK096405 | MK096280 | |
CFCC 52870* | China | Betula platyphylla | MK096321 | MK096361 | NA | MK096406 | MK096281 | |
CFCC 52871* | China | Betula platyphylla | MK096322 | MK096362 | NA | MK096407 | MK096282 | |
CFCC 52872* | China | Betula platyphylla | MK096323 | MK096363 | NA | MK096408 | MK096283 | |
CFCC 52873* | China | Betula platyphylla | MK096350 | MK096390 | NA | MK096435 | MK096310 | |
CFCC 52874* | China | Betula platyphylla | MK096351 | MK096391 | NA | MK096436 | MK096311 | |
CFCC 52875* | China | Betula platyphylla | MK096325 | MK096365 | NA | MK096410 | MK096285 | |
Microascospora fragariae | CBS 118.16 | USA | Fragaria sp. | NR156500 | NA | NA | NA | NA |
CBS 128350 | USA | Rubus sp. | JF514854 | NA | NA | NA | NA | |
1-1 | China | Fragaria ananassa | HM854850 | NA | NA | NA | NA | |
1-2 | China | Fragaria ananassa | HM854849 | NA | NA | NA | NA | |
1-3 | China | Fragaria ananassa | HM854852 | NA | NA | NA | NA | |
Microascospora rubi | MFLU 15-1112 | Italy | Rubus ulmifolia | MF190154 | MF190098 | NA | MF377581 | MF377611 |
MFLU 17-0883 | Italy | Rubus ulmifolia | MF190153 | MF190099 | NA | MF377582 | MF377612 | |
Nakataea oryzae | CBS 243.76 | NA | NA | KM484861 | DQ341498 | NA | NA | NA |
Oblongisporothyrium castanopsidis | ATCC 22470 | Japan | Castanopsis cuspidata | MG591850 | MG591943 | NA | MG592038 | MG976454 |
Ophiodiaporthe cyatheae | YMJ1364 | China | Cyathea lepifera | JX570889 | JX570891 | NA | JX570893 | NA |
Pachytrype princeps | Rogers S | USA | NA | NA | FJ532382 | NA | NA | NA |
Pachytrype rimosa | FF1066 | Costa Rica | NA | NA | FJ532381 | NA | NA | NA |
Paradiaporthe artemisiae | MFLUCC 14-0850 | Italy | Artemisia sp. | MF190155 | MF190100 | NA | NA | NA |
MFLUCC 17-1663 | Italy | Artemisia sp. | MF190156 | MF190101 | NA | NA | NA | |
Phaeoappendispora thailandensis | MFLUCC 13-0161 | Thailand | Quercus sp. | MF190157 | MF190102 | NA | MF377613 | NA |
Phaeodiaporthe appendiculata | CBS 123821 = D77 | Austria | Acer campestre | KF570156 | KF570156 | NA | NA | NA |
CBS 123809 = D76 | Austria | Acer campestre | KF570155 | KF570155 | NA | NA | NA | |
Phragmoporthe conformis | AR 3632 = CBS 109783 | Canada | Alnus rubra | DQ323527 | AF408377 | NA | NA | NA |
Plagiostoma euphorbiae | CBS 340.78 | Netherlands | Euphorbia palustris | EU199198 | AF408382 | NA | DQ368643 | NA |
Plagiostoma salicellum | AR 3455 = CBS 109775 | Austria | Salix sp. | DQ323529 | AF408345 | NA | EU199141 | EU221916 |
Prosopidicola mexicana | CBS 113530 | USA | Prosopis glandulosa | AY720710 | NA | NA | NA | NA |
CBS 113529 | USA | Prosopis glandulosa | AY720709 | KX228354 | NA | NA | NA | |
Pseudomelanconis caryae | CFCC 52110 | China | Carya cathayensis | MG682082 | MG682022 | NA | MG682042 | MG682062 |
CFCC 52111 | China | Carya cathayensis | MG682083 | MG682023 | NA | MG682043 | MG682063 | |
CFCC 52112 | China | Carya cathayensis | MG682084 | MG682024 | NA | MG682044 | MG682064 | |
CFCC 52113 | China | Carya cathayensis | MG682085 | MG682025 | NA | MG682045 | MG682065 | |
Pseudoplagiostoma eucalypti | CBS 124807 | Venezuela | Eucalyptus urophylla | GU973512 | GU973606 | NA | NA | NA |
CBS 116382 | Thailand | Eucalyptus camaldulensis | GU973514 | GU973608 | NA | NA | NA | |
Pseudoplagiostoma oldii | CBS 115722 | Australia | Eucalyptus camaldulensis | GU973535 | GU973610 | NA | NA | NA |
Pseudoplagiostoma variabile | CBS 113067 | Uruguay | Eucalyptus globulus | GU973536 | GU973611 | NA | NA | NA |
Pyricularia grisea | Ina168 | NA | NA | AB026819 | AB026819 | NA | NA | NA |
Racheliella saprophytica | NTCL052-1 | Thailand | Syzygium cumini | KJ021933 | KJ021935 | NA | NA | NA |
Racheliella wingfieldiana | CBS 143669 | South Africa | Syzigium guineense | MG591911 | MG592006 | NA | MG592100 | MG976487 |
Rossmania ukurunduensis | AR 3484 | Russia | Acer ukurunduense | NA | EU683075 | NA | NA | NA |
Saprothyrium thailandense | MFLU 13-0260 | Thailand | Decaying leaf | MF190163 | MF190110 | NA | NA | NA |
Sheathospora cornuta | CFCC 51990* | China | Cornus controversa | MF360006 | MF360008 | NA | MF360002 | MF360004 |
CFCC 51991* | China | Juglans regia | MF360007 | MF360009 | NA | MF360003 | MF360005 | |
Sillia ferruginea | AR 3440 = CBS 126567 | Austria | Corylus avellana | JF681959 | EU683076 | NA | NA | NA |
Sphaerosporithyrium mexicanum | CFNL 2945 | Mexico | Quercus eduardi | MG591896 | MG591990 | NA | MG592083 | MG976473 |
Stegonsporium protopyriforme | CBS 117041 | Austria | Acer pseudoplatanus | NR126119 | EU039992 | NA | NA | NA |
Stegonsporium pyriforme | CBS 124487 | UK | Acer heldreichii | KF570160 | KF570160 | NA | KF570190 | NA |
Stilbospora macrosperma | CBS 121883 | Austria | Carpinus betulus | JX517290 | JX517299 | NA | KF570196 | NA |
CBS 121695 | Netherlands | Carpinus betulus | JX517288 | JX517297 | NA | NA | NA | |
Sydowiella depressula | CBS 813.79 | Switzerland | Rubus sp. | NA | EU683077 | NA | NA | NA |
Sydowiella fenestrans | AR 3777 = CBS 125530 | Russia | Chamerion angustifolium | JF681956 | EU683078 | NA | NA | NA |
Synnemasporella aculeans | AR 3878 = CBS 126566 | USA | Rhus glabra | NA | EU255134 | NA | NA | NA |
CFCC 52094 | China | Rhus chinensis | MG682086 | MG682026 | NA | MG682046 | MG682066 | |
CFCC 52095 | China | Rhus chinensis | MG682087 | MG682027 | NA | MG682047 | MG682067 | |
CFCC 52096 | China | Rhus chinensis | MG682088 | MG682028 | NA | MG682048 | MG682068 | |
Synnemasporella toxicodendri | CFCC 52097 | China | Toxicodendron sylvestre | MG682089 | MG682029 | NA | MG682049 | MG682069 |
CFCC 52098 | China | Toxicodendron sylvestre | MG682090 | MG682030 | NA | MG682050 | MG682070 | |
CFCC 52099 | China | Toxicodendron sylvestre | MG682091 | MG682031 | NA | MG682051 | MG682071 | |
Tubakia japonica | ATCC 22472 | Japan | Castanea crenata | MG591886 | MG591978 | NA | MG592071 | MG976465 |
CBS 191.71 | Japan | Castanea crenata | MG591885 | MG591977 | NA | MG592070 | MG976464 | |
MUCC 2297 | Japan | Castanea crenata | NA | MG591979 | NA | MG592072 | MG976466 | |
MUCC 2298 | Japan | Castanea crenata | NA | MG591980 | NA | MG592073 | MG976467 | |
MUCC 2300 | Japan | Castanea crenata | NA | MG591981 | NA | MG592074 | MG976468 | |
MUCC 2301 | Japan | Castanea crenata | NA | MG591982 | NA | MG592075 | MG976469 | |
Tubakia seoraksanensis | CBS 127490 | South Korea | Quercus mongolica | MG591907 | KP260499 | NA | MG592094 | NA |
Tubakia sutoniana | ICMP 14042 | New Zealand | Quercus sp. | KC145909 | NA | NA | NA | KC145954 |
ICMP 14043 | New Zealand | Quercus ilex | KC145858 | NA | NA | NA | KC145955 |
Species identification was based on morphological features of the ascomata or conidiomata produced on infected plant tissues and micromorphology, supplemented by cultural characteristics. Cross-sections were prepared by hand using a double-edge blade under a dissecting microscope. More than 10 conidiomata/ascomata, 10 asci and/or 50 conidia/ascospores were measured to calculate the mean size and standard deviation (SD). Microscopic photographs were captured with a Nikon Eclipse 80i microscope equipped with a Nikon digital sight DS-Ri2 high definition colour camera, using differential interference contrast (DIC) illumination and the Nikon software NIS-Elements D Package v. 3.00. Adobe Bridge CS v. 6 and Adobe Photoshop CS v. 5 were used for the manual editing. Nomenclatural novelties and descriptions were deposited in MycoBank (
Genomic DNA was extracted using a modified CTAB method, with fungal mycelium harvested from PDA plates with cellophane (
DNA sequences generated by each primer combination were used to obtain consensus sequences using SeqMan v. 7.1.0 in the DNASTAR Lasergene Core Suite software package (DNASTAR Inc., Madison, WI, USA). Reference sequences were selected based on ex-type or ex-epitype sequences available from relevant published literature (
A partition homogeneity test (PHT) with heuristic search and 1 000 search replicates was performed using PAUP to test for incongruence amongst the ITS, LSU, RPB2 and TEF1-α sequence datasets in reconstructing phylogenetic trees. Maximum parsimony (MP) analysis was run using 1 000 heuristic search replicates with random-additions of sequences with a tree bisection and reconnection (TBR) algorithm. Maxtrees were set to 5 000, branches of zero length were collapsed and all equally parsimonious trees were saved. Other calculated parsimony scores were tree length (TL), consistency index (CI), retention index (RI) and rescaled consistency (RC). Maximum likelihood (ML) analysis was performed with a GTR site substitution model, including a gamma-distributed rate heterogeneity and a proportion of invariant sites (
MrModeltest v. 2.3 was used to estimate the best nucleotide substitution model settings for each gene (
In addition to the above analyses, we provided separate phylogenetic trees for Juglanconidaceae, Melanconidaceae and Melanconiellaceae, based on various gene regions (see below) and the same analyses parameters as given above. Phylograms were edited using FigTree v. 1.3.1 (
The combined matrix of ITS, LSU, RPB2 and TEF1-α of Diaporthales included 209 ingroup and two outgroup taxa, comprising 3 269 characters including gaps (776 characters for ITS, 517 for LSU, 1107 for RPB2 and 869 for TEF1-α) in the aligned matrix. Of these, 1 417 characters were constant, 192 variable characters were parsimony-uninformative and 1 660 characters were parsimony informative. The MP analysis resulted in 100 most parsimonious trees (TL = 10 370, CI = 0.341, RI = 0.806, RC = 0.275) and the first tree is shown as Fig.
Phylogram of Diaporthales obtained from an MP analysis of a combined matrix of ITS, LSU, RPB2 and TEF1-α. MP and ML bootstrap support values above 50% are shown at the first and second position, respectively. Thickened branches represent posterior probabilities above 0.95 from BI. Scale bar = 200 changes. Type species are in bold. Strains obtained in the current study are in blue.
For the single genus Juglanconis (Juglanconidaceae), a combined ITS, LSU, CAL and RPB2 matrix of 23 ingroup accessions (five from this study and 18 retrieved from GenBank) was produced, which comprised 2 736 characters including gaps (2 427 constant, 216 variable and parsimony-uninformative, 93 parsimony-informative). A heuristic MP search generated nine equally most parsimonious trees (TL = 332, CI = 0.976, RI = 0.985, RC = 0.961), one of which is shown in Fig.
Phylogram of Juglanconis (Juglanconidaceae) obtained from an MP analysis of a combined matrix of ITS, LSU, CAL and RPB2. MP and ML bootstrap support values above 50% are shown at the first and second position, respectively. Thickened branches represent posterior probabilities above 0.95 from BI. Scale bar = 20 changes. Type species are in bold. Strains obtained in the current study are in blue.
For Melanconiellaceae, a combined ITS, LSU, RPB2 and TEF1-α matrix was produced from 53 ingroup accessions (six from this study and 47 retrieved from GenBank), which comprised 4 122 characters including gaps (2 829 constant, 87 variable and parsimony-uninformative, 1 206 parsimony-informative). A heuristic MP search generated 24 most parsimonious trees (TL = 2 716, CI = 0.652, RI = 0.880, RC = 0.573), one of which is shown in Fig.
For the single genus Melanconis (Melanconidaceae), a combined ITS, LSU, RPB2 and TEF1-α matrix was produced for 57 ingroup accessions (49 from this study and eight retrieved from GenBank), which comprised 2 597 characters including gaps (2 238 constant, 219 variable and parsimony-uninformative, 140 parsimony-informative). A heuristic MP search generated 144 most parsimonious trees (TL = 459, CI = 0.861, RI = 0.919, RC = 0.791), one of which is shown in Fig.
Juglanconis Voglmayr & Jaklitsch, Persoonia 38: 142 (2017)
Juglanconidaceae, with the single genus Juglanconis, was newly introduced by
Juglanconis juglandina (Kunze) Voglmayr & Jaklitsch, Persoonia 38: 144 (2017).
Juglanconis was newly introduced by
≡Melanconium juglandinum Kunze, Fl. Dresd., 2. Aufl.: 260. 1823.
Conidiomata acervular, immersed in host bark, erumpent from surface of host branches, scattered or occasionally confluent, 1.5–2.5 mm, covered by black discharged conidial masses at maturity, usually conspicuous. Ectostromatic disc straw to honey, surrounded by bark or not. Central column beneath the disc more or less conical, straw to buff. Conidiophores cylindrical to lageniform, simple, rarely branched at the base, smooth, subhyaline to pale brown. Conidiogenous cells annellidic with distinct annellations, integrated. Conidia unicellular, initially hyaline, becoming brown to blackish when mature, broadly ellipsoid to broadly pip-shaped, truncate with distinct scar at the base, densely multiguttulate, thick-walled, (17–)19–22(–24.5) × (9–)11–14(–16.5) μm (av. = 20 × 13 μm, n = 50), with 0.8–1 µm wide gelatinous sheath. Sexual morph was not observed.
On PDA, cultures are initially white, becoming straw after 3–5 d and grey olivaceous after 7–10 d. The colonies are felty with an irregular edge; sterile.
(all on twigs and branches of Juglans regia). CHINA, Gansu Province, Qingyang City, Shishe village, 35°38'17.08"N, 107°47'48.68"E, 14 July 2013, X.L. Fan (BJFC-S908; living culture, CFCC 51727); Gansu Province, Qingyang City, Zhongwan Forest Farm, 35°26'26.33"N, 108°34'09.38"E, 11 July 2013, X.L. Fan (BJFC-S947; living culture, CFCC 51728); Gansu Province, Qingyang City, Zhongwan Forest Farm, 35°26'25.52"N, 108°34'09.03"E, 11 July 2013, X.L. Fan (BJFC-S955; living culture, CFCC 51729).
Juglanconis juglandina is the type species of Juglanconis and is thus far only known to occur on Juglans regia distributed in Asia and Europe (
≡ Melanconium oblongum Berk., Grevillea 2 (no. 22): 153. 1874.
= Diaporthe juglandis Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 45: 448. 1893.
≡ Melanconis juglandis (Ellis & Everh.) A.H. Graves, Phytopathology 13: 311. 1923.
Pseudostromata immersed in host bark, distinctly erumpent from surface of host branches, 1.5–3 mm diam. Ectostromatic disc indistinct, usually circular, greyish to brownish. Perithecia often appearing as rounded bumps beneath the bark surface surrounding the ectostromatic disc, prolonged black neck from the top, (450–)525–700(–780) µm diam. (av. = 580 μm, n = 30). Asci hyaline, clavate to fusoid, (120–)122–135 × (12.5–)13–16.5 (–17) μm (av. = 126.5 × 15 μm, n = 20). Ascospores hyaline, ellipsoid, broadly ellipsoid or broadly fusoid, symmetric to slightly asymmetric, straight, rarely slightly curved, constricted at the septum, (17–)17.5–22(–23.5) × (7.5–)8–10.5(–11) μm (av. = 19.5 × 9.5 μm, n = 50). Conidiomata acervular, immersed in host bark, erumpent from surface of host branches, scattered or occasionally confluent, 1–2 mm, covered by black discharged conidial masses at maturity, usually conspicuous. Ectostromatic disc buff to honey, surrounded by bark or not. Central column beneath the disc more or less conical, isabelline to olivaceous grey. Conidiophores cylindrical to lageniform, simple, rarely branched at the base, smooth, subhyaline to pale brown. Conidiogenous cells annellidic with distinct annellations, integrated. Conidia unicellular, initially hyaline, becoming brown to blackish when mature, broadly ellipsoid to broadly pip-shaped, truncate with distinct scar at the base, densely multiguttulate, thick-walled, (14–)19–23.5(–28) × (6.5–)9–13(–15) μm (av. = 22 × 12.5 μm, n = 50), with 0.8–1 µm wide gelatinous sheath.
Morphology of Juglanconis oblonga from Juglans regia. A–B habit of acervuli on branches C transverse section through acervulus D longitudinal section through perithecia E longitudinal section through acervulus F conidiophores, conidiogenous cells G conidia H asci and ascospores I ascospores. Scale bars: 10 mm (A), 500 μm (B–E), 20 μm (F–I).
On PDA, cultures are initially white, becoming pale olivaceous grey after 10 d. The colonies are felty with an irregular edge; texture uniform; sterile.
(all on twigs and branches of Juglans regia). CHINA, Heilongjiang Province, Harbin City, Linan, Heilongjiang Botanical Garden, 45°42'21.10"N, 126°38'42.87"E, 2 August 2016, Q. Yang & Z. Du (BJFC-S1374; living culture, CFCC 51725; ibid.CFCC 51726).
Juglanconis oblonga is based on Melanconium oblongum (= Melanconis juglandis) (
Melanconis Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 115 (1863)
Melanconidaceae was introduced by Winter (1886) and subsequently involved many genera with perithecia immersed in a well-developed stroma with ostioles (beaks) that emerge through an ectostromatic disc (
Phylogram of Melanconis (Melanconidaceae) obtained from an MP analysis of a combined matrix of ITS, LSU, RPB2 and TEF1-α. MP and ML bootstrap support values above 50% are shown at the first and second position, respectively. Thickened branches represent posterior probabilities above 0.95 from BI. Scale bar = 20 changes. Type species are in bold. Strains obtained in the current study are in blue.
Melanconis stilbostoma (Fr.) Tul. & C. Tul., Select. fung. carpol. (Paris) 2: 115 (1863)
The type genus Melanconis was established by Tulasne and Tulasne (1863) based on Sphaeria stilbostoma Fr. This genus is characterised by circularly arranged perithecia immersed in well developed to reduced entostromata with a concolourous central column and ostioles erumpent through a light-coloured ectostromatic disc with hyaline, one-septate ascospores; acervuli with light-coloured central column producing brown to olive-brown, fusiform to pyriform alpha conidia and hyaline, cylindrical or allantoid beta conidia (
(all on twigs and branches of Betula albosinensis). CHINA, Gansu Province, Gannan Tibetan Autonomous Prefecture, Zhouqu County, Qiban Forestry Centre, 33°56'35.36"N, 104°07'13.03"E, 20 August 2014, Y.M. Liang (BJFC-S1319, holotype; living ex-type culture, CFCC 50471); Gansu Province, Gannan Tibetan Autonomous Prefecture, Zhouqu County, Qiban Forestry Centre, 33°56'37.05"N, 104°07'13.78"E, 20 August 2014, Y.M. Liang (BJFC-S13200; living culture, CFCC 50472); Gansu Province, Gannan Tibetan Autonomous Prefecture, Zhouqu County, Qiban Forestry Centre, 33°56'34.44"N, 104°07'15.59"E, 20 August 2014, Y.M. Liang (BJFC-S1321; living culture, CFCC 50473).
Melanconis betulae was described from Betula albosinensis (
(all on twigs and branches of Betula albosinensis). CHINA, Gansu Province, Gannan Tibetan Autonomous Prefecture, Zhouqu County, Qiban Forestry Centre, 33°56'34.49"N, 104°07'15.21"E, 20 August 2014, X.L. Fan (BJFC-S1322; living culture, CFCC 50474); Shaanxi Province, Ankang City, Ningshan County, Huoditang Forest Farm, 33°26'24.80"N, 108°26'45.10"E, 3 August 2015, Q. Yang (BJFC-S1349; living culture, CFCC 52877; ibid, CFCC 52878); Jilin Province, Jiaohe City, Haiqing Forest Farm, 43°79'88.71"N, 127°15'83.04"E, 26 June 2017, X.W. Wang (CF 20170668; living culture, CFCC 52876).
Melanconis itoana was described from Betula ermanii in Japan (
(all on twigs and branches of Betula platyphylla). CHINA, Tibet Autonomous Region, Linzhi City, Juemu Valley, 29°39'50.13"N, 94°18'50.70"E, 22 July 2016, X.L. Fan (CF 20160703; living culture, CFCC 528433); Heilongjiang Province, Yichun City, Dailing District, Liangshui Natural Reserve, 47°11'05.26"N, 128°57'26.15"E, 29 July 2016, Q. Yang & Z. Du (CF 20161703; living culture, CFCC 52867); Heilongjiang Province, Harbin City, Heilongjiang Botanical Garden, 45°42'27.58"N, 126°38'36.72"E, 2 August 2016, Q. Yang & Z. Du (CF 20161709; living culture, CFCC 52868); Qinghai Province, Menyuan City, Xianmi Forest Farm, 37°16'35.27"N, 101°46'53.78"E, 3 September 2016, J.H. Zuo (CF 20160911; living culture, CFCC 52865); Ningxia Autonomous Region, Yinchuan City, Helan County, Taihedizhonghai, 38°31'50.40"N, 106°17'46.10"E, 5 August 2015, X.L. Fan & Z. Du (CF 20150802; living culture, CFCC 52873); Ningxia Autonomous Region, Jingyuan City, Jingguan Road, 35°29'50.32"N, 106°18'27.10"E, 13 August 2014, X.L. Fan & Z. Du (BJFC-S1324; living culture, CFCC 50476); Beijing City, Tongzhou District, Song Village, 35°59'49.50"N, 116°39'32.35"E, 20 May 2015, X.L. Fan (BJFC-S1325; living culture, CFCC 50477); other materials with similar locations and hosts are listed in Table
Melanconis stilbostoma is the type species of Melanconis and is thus far only known to occur on Betula spp. with a worldwide distribution (
Melanconiella Sacc., Syll. fung. (Abellini) 1: 740 (1882)
Melanconiellaceae was validated by
Phylogram of Melanconiellaceae obtained from an MP analysis from a combined matrix of ITS, LSU, RPB2 and TEF1-α. MP and ML bootstrap support values above 50% are shown at the first and second position, respectively. Thickened branches represent posterior probabilities above 0.95 from BI. Scale bar = 80 changes. Type species are in bold. Strains obtained in the current study are in blue.
Melanconiella spodiaea (Tul. & C. Tul.) Sacc., Syll. fung. (Abellini) 1: 740 (1882)
The genus Melanconiella was established by
betulicola (Lat.): referring to the host genus on which it was collected, Betula.
This species is distinguished by hyaline ascospores, (16.5–)18–22(–24) × (3–)4–6 μm, with slightly constricted at the septum and with hyaline broad cap-like appendages at both ends.
CHINA. Shaanxi Province: Ningshan County, Huoditang Forest Farm, Huodi Valley, 33°26'36.32"N, 108°26'46.48"E, 3 August 2015, on twigs and branches of Betula albosinensis, Q. Yang (BJFC-S1347 holotype; living culture, CFCC 52482).
Pseudostromata inconspicuous, immersed in host bark, slightly erumpent from surface of host branches, 1.5–3 mm diam. Ectostromatic disc indistinct, usually circular, buff to hazel. Central column circular, mouse grey to iron grey. Ostioles numerous, violaceous black to black, scarcely projecting, 70–150 μm diam. Perithecia flask-shaped to spherical, arranged circularly or irregularly, 7–12 per disc, often appearing as rounded bumps beneath the bark surface surrounding the ectostromatic disc, (320–)350–550(–610) µm diam. (av. = 480 μm, n = 30). Asci hyaline, clavate to fusoid, (50–)55–65(–70) × (7–)8.5–14(–16) μm (av. = 60 × 11 μm, n = 20). Ascospores hyaline, ellipsoid, broadly ellipsoid or broadly fusoid, 2–4 guttulate, symmetric to slightly asymmetric, straight, rarely slightly curved, slightly constricted at the septum, (16.5–)18–22(–24) × (3–)4–6 μm (av. = 20 × 4.5 μm, n = 50), with hyaline broad cap-like appendages at both ends. Conidiomata acervular, immersed in host bark, erumpent from surface of host branches, scattered or occasionally confluent, 1.3–2.5 mm, covered by fawn to dark brick discharged conidial masses at maturity, usually conspicuous. Ectostromatic disc inconspicuous. Central column beneath the disc more or less conical, olivaceous grey to iron grey. Conidiophores hyaline, smooth, cylindrical to lageniform, simple, rarely branched at the base. Conidiogenous cells hyaline, phialidic. Conidia unicellular, hyaline, narrowly ellipsoid, elongate to slightly allantoid, (9.5–)10–13.5(–15) × (2–)3–4.5(–5.5) μm (av. = 13 × 3.5 µm, n = 50), with 0.5 µm wide gelatinous sheath.
Morphology of Melanconiella betulicola from Betula albosinensis. A–B habit of pseudostromata on branches C transverse section through perithecia D longitudinal section through perithecia E–F habit of acervuli on branches G transverse section through acervulus H longitudinal section through acervulus I asci and ascospores J–K ascus and ascospores L–O ascospores P conidiophores, conidiogenous cells and conidia Q conidia. Scale bars: 2 mm (A, E), 500 μm (B–D, F–H), 10 μm (J–K, P–Q), 5 μm (L–O).
On PDA, cultures are initially white, becoming greyish-sepia after 3 d and distensible radially after 10 d. The colonies are felty with an irregular edge; texture uniform; sterile.
CHINA. Shaanxi Province: Ningshan County, Huoditang Forest Farm, Huodi Valley, 33°26'37.53"N, 108°26'44.14"E, 3 August 2015, on twigs and branches of Betula albosinensis, Q. Yang (CF 20150847; living culture, CFCC 52483);
Melanconiella betulicola is associated with canker disease of Betula albosinensis in China. It is similar to M. ellisii but differs by larger ascospores (18–22 × 4–6 vs.> 12.5–16 × 4.0–5.5 μm) with hyaline, broad cap-like appendages at both ends (
corylina (Lat.): referring to the host genus on which it was collected, Corylus.
This species is distinguished by acervuli erumpent through circularly cracked host bark and covered by olivaceous buff to honey discharged conidial masses at maturity; conidia unicellular, hyaline, with various shapes and 1–3 guttulate, (7–)8–13.5(–14.5) × (2–)2.5–4(–5) μm.
Morphology of Melanconiella corylina from Corylus mandshurica. A habit of acervuli on branches B–F process of development of acervulus G transverse section through acervulus H–I longitudinal section through acervulus J conidiophores K conidiogenous cells and conidia L–W conidia. Scale bars: 2 mm (A), 500 μm (B–I), 10 μm (J–K), 5 μm (L–W).
CHINA. Shaanxi Province: Baoji County, Taibai Mountain, 34°15'43.32"N, 107°88'42.16"E, 13 July 2017, on twigs and branches of Corylus mandshurica, N. Jiang (BJFC-FB56 holotype; living culture, CFCC 52484).
Conidiomata acervular, immersed in host bark, erumpent from surface of host branches, scattered or occasionally confluent, 1–1.5 mm, erumpent through circularly cracked host bark and covered by olivaceous buff to honey discharged conidial masses at maturity, usually conspicuous. Ectostromatic disc inconspicuous and cracked circularly at maturity. Central column beneath the disc more or less oblate, iron grey to dark grey. Conidiophores hyaline, smooth, cylindrical, simple, rarely branched at the base. Conidiogenous cells hyaline, phialidic. Conidia unicellular, hyaline, narrowly ellipsoid to fusoid, elongate to slightly allantoid, 1–3 guttulate, (7–)8–13.5(–14.5) × (2–)2.5–4(–5) μm (av. = 10 × 3.5 µm, n = 50) μm (av. = 13 × 3.5 µm, n = 50). Sexual morph was not observed.
On PDA, cultures are initially white, becoming fuscous black in the centre and edge after 5 d. The colonies are felty with an irregular edge; texture uniform; sterile.
CHINA. Shaanxi Province: Baoji County, Taibai Mountain, 34°15'40.05"N, 107°88'43.33"E, 13 July 2017, on twigs and branches of Corylus mandshurica, N. Jiang (CF 20170756 holotype; living culture, CFCC 52485).
Melanconiella corylina is associated with canker disease of Corylus mandshurica in China. It can be distinguished from its closest relative, the generic type M. spodiaea growing in Carpinus spp., by its hyaline, discosporina-like conidia, and the smaller size of conidia (8–13.5 × 2.5–4 vs.> 13.3–15.2 × 7.5–8.5 μm) as well as the hosts (
Sheathospora (Lat.): referring to the conidia with distinct hyaline sheath.
This genus differs from other genera in Melanconiellaceae by conical and discrete pycnidia with aseptate, cylindrical to ellipsoidal conidia with distinct hyaline sheath.
Sheathospora cornuta (C.M. Tian & Z. Du) Fan.
Conidiomata pycnidial, immersed in host bark, erumpent through the surface of host branches. Ectostromatic disc inconspicuous and extended to form a beak at maturity. Central column absent. Conidiophores hyaline, smooth, cylindrical, simple, rarely branched at the base. Conidiogenous cells hyaline, phialidic. Conidia hyaline, aseptate, with distinct hyaline sheath. Sexual morph was not observed.
Sheathospora is established for Melanconiella cornuta, which was previously included in the Melanconiella clade (
Melanconiella cornuta C.M. Tian & Z. Du, Phytotaxa 327(3): 257 (2017)
This species is distinguished by conical and discrete pycnidia without central column and aseptate, cylindrical to ellipsoidal, (19–)19.5–22.5(–23) × (8–)8.5–10.5(–11) μm conidia, with a distinct hyaline sheath 1–1.5 μm wide.
CHINA. Shaanxi Province: Ankang City, Ningshan County, Huoditang Forest Farm, 33°26'04.46"N, 108°26'59.91"E, 3 July 2016, on twigs and branches of Cornus controversa, X.L. Fan (BJFC-S1375 holotype; living ex-type culture CFCC 51990).
Conidiomata pycnidial, immersed in host bark, conical, with single necks erumpent through the surface of host branches, scattered, (250–)270–330(–410) μm (av. = 300 μm, n = 20) diam. Ectostromatic disc inconspicuous and extended to form a beak at maturity, pale luteous to amber. Central column absent. Conidiophores hyaline, smooth, cylindrical, simple, rarely branched at the base, 17–24(–25) × 2.5–4(–4.5) μm (av. = 21.5 × 3.5 µm, n = 50). Conidiogenous cells hyaline, phialidic. Conidia hyaline, aseptate, cylindrical to ellipsoidal, (19–)19.5–22.5(–23) × (8–)8.5–10.5(–11) μm (av. = 21 × 10 µm, n = 50), with distinct hyaline sheath, 1–1.5 μm wide at maturity. Sexual morph was not observed.
Colony growth on PDA originally white, becoming pale yellowish after 7–10 days. Colony flat, felty-like, with a uniform texture and yellowish to dark brown conidiomata irregularly scattered on the colony surface.
CHINA. Shaanxi Province: Ankang City, Ningshan County, Huoditang Forest Farm, 36°26'13.30"N, 108°26'48.32"E, 3 August 2015, on twigs and branches of Juglans regia, Q. Yang (BJFC-S1345 paratype; living ex-paratype culture CFCC 51991).
Sheathospora cornuta is proposed here as a new combination for Melanconiella cornuta. It is the type and currently only species of Sheathospora and so far known from Cornus controversa and Juglans regia in China. The sexual morph of this species is unknown and further collections are required to elucidate its life cycle.
During the investigation of melanconis-like fungi in China, we identified eight species residing in three families (Juglanconidaceae, Melanconidaceae and Melanconiellaceae) of Diaporthales. It includes Juglanconis juglandina, J. oblonga, Melanconis betulae, Ms. itoana, Ms. stilbostoma, the two new species Melanconiella betulicola and M. corylina and the new combination Sheathospora cornuta in the new genus Sheathospora.
All specimens in the current study were collected from symptomatic branches and twigs associated with canker or dieback diseases, of which Juglanconis (Juglanconidaceae) species were isolated from Juglans regia (Juglandaceae), Melanconiella (Melanconiellaceae) species from Betula albosinensis and Corylus mandshurica (Betulaceae) and Melanconis (Melanconidaceae) species from Betula albosinensis and Betula platyphylla (Betulaceae). It may indicate that many melanconis-like species have obvious host specificity. The type species of the new genus Sheathospora (Melanconiellaceae) was isolated from Cornaceae (Cornus controversa) and Juglans regia (Juglandaceae), suggesting a low host specificity and that additional undiscovered hosts species of this taxon may exist in China.
As the morphological features in previous melanconis-like fungi are highly overlapping, phylogenetic studies using DNA sequences have been useful to elucidate the diversity and systematics in this group. The current results indicated that Juglanconis and Melanconis are still unique, the only genera in Juglanconidaceae and Melanconidaceae, respectively, due to the lacking of extensive fresh collections. The family Melanconiellaceae was recently proposed by
As shown in this paper, future studies addressing the fungal diversity associated with canker or dieback diseases should routinely include sequence data for protein-coding genes to achieve stable, supported topologies in phylogenetic trees. It is hoped that the classification proposed here will also provide an updated phylogenetic framework that will facilitate further revision of the families with melanconis-like asexual morphs. Although the current study provides additional new data on melanconis-like genera, typification, species concept and taxonomic affiliation of many described Melanconium species are yet unclear, including the type species M. atrum, which currently represents a doubtful taxon (
This study is financed by National Natural Science Foundation of China (Project No.: 31670647) and National Key R&D Program of China (Project No.: 2017YFD0600105). All authors want to thank the Experimental Teaching Centre (College of Forestry, Beijing Forestry University) for providing installed scientific equipments during the whole process.