Research Article |
Corresponding author: Lei Cai ( cail@im.ac.cn ) Academic editor: Paul Cannon
© 2018 Mei Wang, Xiao-Ming Tan, Fang Liu, Lei Cai.
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:
Wang M, Tan X-M, Liu F, Cai L (2018) Eight new Arthrinium species from China. MycoKeys 34: 1-24. https://doi.org/10.3897/mycokeys.34.24221
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The genus Arthrinium includes important plant pathogens, endophytes and saprobes with a wide host range and geographic distribution. In this paper, 74 Arthrinium strains isolated from various substrates such as bamboo leaves, tea plants, soil and air from karst caves in China were examined using a multi-locus phylogeny based on a combined dataset of ITS rDNA, TEF1 and TUB2, in conjunction with morphological characters, host association and ecological distribution. Eight new species were described based on their distinct phylogenetic relationships and morphological characters. Our results indicated a high species diversity of Arthrinium with wide host ranges, amongst which, Poaceae and Cyperaceae were the major host plant families of Arthrinium species.
Ascomycota , Morphology, Phylogeny, Systematics, Taxonomy
Arthrinium Kunze is an anamorph-typified genus, which has been traditionally linked to the teleomorph-typified genus Apiospora Sacc. (
Arthrinium species are geographically widely distributed in various hosts. Many species of Arthrinium are associated with plants as endophytes or saprobes, as well as plant pathogens on some important ornamentals, e.g. A. phaeospermum causing culm rot on Phyllostachys viridis (
In this paper, eight new Arthrinium species are described and characterised based on morphological characters and phylogeny inferred from the combined ITS rDNA, TEF1 and TUB2 sequences dataset. Comparisons were made with morphologically similar and phylogenetically related species. Fungus-host distribution of Arthrinium species are summarised based on data from literature and this study.
Diseased and healthy tissues of bamboo leaves and other plant hosts were collected from six provinces or municipalities in China (Chongqing, Guangxi, Guangdong, Guizhou, Jiangxi, Hunan). Tissue pieces (5 mm × 5 mm) were taken from the margin of leaf lesions and the surface sterilised with 75% ethanol for 1 min, 5% NaClO for 30 s, followed by rinsing in sterile distilled water for 1 min. The pieces were dried with sterilised paper towels and then placed on 1/4 PDA (potato dextrose agar) (
All cultures were preserved in the LC culture collection (personal culture collection of Lei Cai housed in the Institute of Microbiology, Chinese Academy of Sciences). Type specimens were deposited in Mycological Herbarium of the Institute of Microbiology, Chinese Academy of Sciences, Beijing, China (
Cultures were incubated on PDA for 7 d at 25 °C to measure the growth rates and on 2% malt agar with bamboo leaves to enhance sporulation. Morphological descriptions were based on cultures sporulating on MEA (malt extract agar) medium at room temperature (ca. 25 °C). Shape and size of microscopic structures were observed using a light microscope and colonies were assessed according to the colour charts of
Fresh fungal mycelia were taken from 7-d-old cultures growing on PDA and ground with the organisation disruptor FastPrep-48. Genomic DNA was extracted following the modified CTAB protocol as described in
Phylogenetic analyses were conducted using partial sequences of three loci, 5.8S nuclear ribosomal gene with the two flanking transcribed spacers (ITS), part of the translation elongation factor 1-alpha (TEF1) and beta-tubulin (TUB2). The ITS locus was amplified using the primer pair ITS1/ITS4 (
PCR was performed in a 25 ml reaction containing 18.95 µl double distilled water, 2.5 µl 10 × PCR buffer, 0.3 µl dNTP mix (2.5 mM), 1 µl of each primer (10 mM), 1 µl DNA template and 0.25 µl Taq DNA polymerase (Genstar). The annealing temperatures were adjusted to 52 °C for ITS and TUB2, and 56 °C for TEF1. Purification and sequencing of the PCR amplicons were done by SinoGenoMax, Beijing.
Sequences generated from the forward and reverse primers were used to obtain consensus sequences using MEGA v. 6.0 (
Bayesian analysis was conducted using MrBayes v. 3.2.1 (
Phylogenetic tree based on the combined ITS, TEF1 and TUB2 sequences alignment generated from a Maximum likelihood phylogenetic analysis. Bootstrap support values (>70%) and posterior probabilities (>0.9) are given at the nodes (ML/PP). The tree is rooted with Nigrospora gorlenkoana
Speices | Strain numbers1 | Hosts | Countries | GenBank accessions | ||
---|---|---|---|---|---|---|
ITS | TUB | TEF | ||||
Arthirinium arundinis |
|
Leaf of Hordeum vulgare | Iran | KF144884 | KF144974 | KF145016 |
|
Living leaves of Fagus sylvatica | Switzerland | KF144885 | KF144975 | KF145017 | |
LC4477 | Unknow host | China | KY494688 | KY705159 | KY705087 | |
LC4493 | Phyllostachys sp. | China | KY494689 | KY806202 | KY705088 | |
LC4650 | Osmanthus sp. | China | KY494695 | KY705165 | KY705094 | |
LC4951 | Dichotomanthus tristaniaecarpa | China | KY494698 | KY705168 | KY705097 | |
LC4959 | Bothrocaryum controversum | China | KY494699 | KY705169 | KY705098 | |
LC5311 | Air in karst cave | China | KY494706 | KY705175 | KY705105 | |
LC5312 | Air in karst cave | China | KY494707 | KY705176 | KY705106 | |
LC5332 | Air in karst cave | China | KY494710 | KY705179 | KY705109 | |
LC5394 | Soil in karst cave | China | KY494711 | KY705180 | KY705110 | |
LC5416 | Water in karst cave | China | KY494712 | KY705181 | KY705111 | |
LC7118 | Leaf of bamboo | China | KY494723 | KY705191 | KY705120 | |
LC7122 | Leaf of bamboo | China | KY494726 | KY705194 | KY705123 | |
LC7160 | Leaf of bamboo | China | KY494738 | KY705206 | KY705134 | |
LC7211 | Leaf of bamboo | China | KY494739 | KY705207 | KY705135 | |
LC7216 | Leaf of bamboo | China | KY494741 | KY705209 | KY705137 | |
LC7218 | Leaf of bamboo | China | KY494742 | KY705210 | KY705138 | |
LC7243 | Leaf of bamboo | China | KY494744 | KY705212 | KY705140 | |
LC7252 | Leaf of bamboo | China | KY494747 | KY705215 | KY705143 | |
LC7277 | Leaf of bamboo | China | KY494750 | KY705218 | KY705146 | |
A. aureum |
|
Air | Spain | AB220251 | KF144981 | KF145023 |
A. bambusae |
LC7106* = |
Leaf of bamboo | China | KY494718 | KY705186 | KY806204 |
LC7107 | Leaf of bamboo | China | KY494719 | KY705187 | KY705117 | |
LC7113 | Leaf of bamboo | China | KY494720 | KY705188 | KY806205 | |
LC7124 | Leaf of bamboo | China | KY494727 | KY705195 | KY806206 | |
LC7125 | Leaf of bamboo | China | KY494728 | KY705196 | KY705124 | |
LC7128 | Leaf of bamboo | China | KY494730 | KY705198 | KY705126 | |
LC7246 | Leaf of bamboo | China | KY494745 | KY705213 | KY705141 | |
A. camelliae-sinensis |
LC5007* = |
Camellia sinensis | China | KY494704 | KY705173 | KY705103 |
LC8181 | Brassica capestris | China | KY494761 | KY705229 | KY705157 | |
A. dichotomanthi |
LC4950* = |
Dichotomanthus tristaniaecarpa | China | KY494697 | KY705167 | KY705096 |
LC8175 | Dichotomanthus tristaniaecarpa | China | KY494755 | KY705223 | KY705151 | |
LC8176 | Dichotomanthus tristaniaecarpa | China | KY494756 | KY705224 | KY705152 | |
A. euphorbiae |
|
Bambusa sp. | Bangladesh | AB220241 | AB220288 | – |
A. guizhouense | LC5318 | Air in karst cave | China | KY494708 | KY705177 | KY705107 |
LC5322* =CGMCC3.18334 | Air in karst cave | China | KY494709 | KY705178 | KY705108 | |
A. gutiae |
|
Gut of a grasshopper | India | KR011352 | KR011350 | KR011351 |
A. hispanicum |
|
Maritime sand | Spain | AB220242 | AB220289 | – |
A. hydei |
|
Culms of Bambusa tuldoides | Hong Kong | KF144890 | KF144982 | KF145024 |
LC7103 | Leaf of bamboo | China | KY494715 | KY705183 | KY705114 | |
LC7105 | Leaf of bamboo | China | KY494717 | KY705185 | KY705116 | |
A. hyphopodii |
|
Culms of Bambusa tuldoides | Thailand | KR069110 | – | – |
A. japonicum |
|
Carex despalata (dead leaf) | Japan | AB220262 | AB220309 | – |
|
Carex despalata (leaf) | Japan | AB220264 | AB220311 | – | |
A. garethjonesii | KUMCC 16-0202 | Dead culms of bamboo | China | KY356086 | – | – |
A. jatrophae | MMI 00052* = |
Healthy petiole of Jatropha podagrica | India | JQ246355 | – | – |
A. jiangxiense | LC2831 | Leaf of bamboo | China | KY494686 | KY80620106201 | KY705085 |
LC4494 | Phyllostachys sp. | China | KY494690 | KY705160 | KY705089 | |
LC4541 | Maesa sp. | China | KY494691 | KY705161 | KY705090 | |
LC4547 | Machilus sp. | China | KY494692 | KY705162 | KY705091 | |
LC4577* = |
Maesa sp. | China | KY494693 | KY705163 | KY705092 | |
LC4578 | Camellia sinensis | China | KY494694 | KY705164 | KY705093 | |
LC4993 | Phyllostachys sp. | China | KY494700 | KY806203 | KY705099 | |
LC4997 | Phyllostachys sp. | China | KY494701 | KY705170 | KY705100 | |
LC5001 | Phyllostachys sp. | China | KY494702 | KY705171 | KY705101 | |
LC5004 | Phyllostachys sp. | China | KY494703 | KY705172 | KY705102 | |
LC5015 | Imperata cylindrica | China | KY494705 | KY705174 | KY705104 | |
A. jiangxiense | LC7104 | Leaf of bamboo | China | KY494716 | KY705184 | KY705115 |
LC7154 | Leaf of bamboo | China | KY494736 | KY705204 | KY705132 | |
LC7156 | Leaf of bamboo | China | KY494737 | KY705205 | KY705133 | |
LC7275 | Leaf of bamboo | China | KY494749 | KY705217 | KY705145 | |
A. kogelbergense |
|
Dead culms of Restionaceae | South Africa | KF144892 | KF144984 | KF145026 |
A. longistromum |
|
Decaying bamboo culms | Thailand | KU940141 | – | – |
|
Decaying bamboo culms | Thailand | KU940142 | – | – | |
A. malaysianum |
|
Macaranga hullettii stem colonised by ants | Malaysia | KF144896 | KF144988 | KF145030 |
A. marii |
|
Air | Spain | AB220252 | KF144993 | KF145035 |
A. mediterranei |
|
Air | Spain | AB220243 | AB220290 | – |
A. mytilomorphum |
|
Dead blades of Andropogon sp. | India | KY494685 | – | – |
A. neosubglobosa | JHB006 | Dead culms of bamboo | China | KY356089 | – | – |
KUMCC 16-0203 | Dead culms of bamboo | China | KY356090 | – | – | |
A. obovatum |
LC4940* = |
Lithocarpus sp. | China | KY494696 | KY705166 | KY705095 |
LC8177 | Lithocarpus sp. | China | KY494757 | KY705225 | KY705153 | |
LC8178 | Lithocarpus sp. | China | KY494758 | KY705226 | KY705154 | |
A. ovatum |
|
Arundinaria hindsii | Hong Kong | KF144903 | KF144995 | KF145037 |
A. paraphaeospermum | MFLU 16-1974 | Dead clumps of Bambusa sp. | Thailand | KX822128 | – | – |
A. phaeospermum |
|
Leaf of Hordeum vulgare | Iran | KF144904 | KF144996 | KF145038 |
|
Leaf of Hordeum vulgare | Iran | KF144905 | KF144997 | KF145039 | |
|
Leaf of Hordeum vulgare | Iran | KF144906 | KF144998 | KF145040 | |
|
Leaf of Hordeum vulgare | Iran | KF144907 | KF144999 | KF145041 | |
A. phragmites | CPC18900* | Culms of Phragmites australis | Italy | KF144909 | KF145001 | KF145043 |
A. pseudoparenchymaticum |
LC7234* = |
Leaf of bamboo | China | KY494743 | KY705211 | KY705139 |
LC8173 | Leaf of bamboo | China | KY494753 | KY705221 | KY705149 | |
LC8174 | Leaf of bamboo | China | KY494754 | KY705222 | KY705150 | |
A. pseudosinense |
|
Leaf of bamboo | The Netherlands | KF144910 | – | KF145044 |
A. pseudospegazzinii |
|
Macaranga hullettii stem colonised by ants | Malaysia | KF144911 | KF145002 | KF145045 |
A. pterospermum |
|
Leaf lesion of Machaerina sinclairii | Australia | KF144913 | KF145004 | KF145046 |
A. puccinioides |
|
Leaf of Lepidosperma gladiatum | Germany | AB220253 | AB220300 | – |
A. rasikravindrii |
|
Cissus sp. | The Netherlands | KF144914 | – | – |
|
Rice | Thailand | KF144915 | – | – | |
|
Dead bamboo culms | Thailand | KU940143 | – | – | |
|
Dead bamboo culms | Thailand | KU940144 | – | – | |
|
Soil | Svalbard | JF326454 | – | – | |
LC5449 | Soil in karst cave | China | KY494713 | KY705182 | KY705112 | |
LC7115 | Leaf of bamboo | China | KY494721 | KY705189 | KY705118 | |
LC7117 | Leaf of bamboo | China | KY494722 | KY705190 | KY705119 | |
LC7119 | Leaf of bamboo | China | KY494724 | KY705192 | KY705121 | |
LC7120 | Leaf of bamboo | China | KY494725 | KY705193 | KY705122 | |
LC7126 | Leaf of bamboo | China | KY494729 | KY705197 | KY705125 | |
LC7129 | Leaf of bamboo | China | KY494731 | KY705199 | KY705127 | |
LC7135 | Leaf of bamboo | China | KY494732 | KY705200 | KY705128 | |
LC7139 | Leaf of bamboo | China | KY494733 | KY705201 | KY705129 | |
LC7141 | Leaf of bamboo | China | KY494734 | KY705202 | KY705130 | |
LC7142 | Leaf of bamboo | China | KY494735 | KY705203 | KY705131 | |
LC7251 | Leaf of bamboo | China | KY494746 | KY705214 | KY705142 | |
LC7254 | Leaf of bamboo | China | KY494748 | KY705216 | KY705144 | |
LC8179 | Brassica capestris | China | KY494759 | KY705227 | KY705155 | |
LC8180 | Brassica capestris | China | KY494760 | KY705228 | KY705156 | |
A. sacchari |
|
Phragmites australis | United Kingdom | KF144916 | KF145005 | KF145047 |
|
Bamboo | Indonesia | KF144917 | KF145006 | KF145048 | |
A. saccharicola |
|
Air | The Netherlands | KF144920 | KF145009 | KF145051 |
|
Dead culms of Phragmites australis | France | AB220257 | KF145010 | KF145052 | |
|
Dead culms of Phragmites australis | The Netherlands | KF144921 | KF145011 | KF145053 | |
A. serenense |
|
Food, pharmaceutical excipients, atmosphere and home dust | Spain | AB220250 | AB220297 | – |
A. subglobosum |
|
Dead bamboo culms | Thailand | KR069112 | – | – |
A. subroseum | LC7215 | Leaf of bamboo | China | KY494740 | KY705208 | KY705136 |
LC7291 | Leaf of bamboo | China | KY494751 | KY705219 | KY705147 | |
LC7292* =CGMCC3.18337 | Leaf of bamboo | China | KY494752 | KY705220 | KY705148 | |
A. thailandicum |
|
Dead bamboo culms | Thailand | KU940145 | – | – |
LC5630 | Rotten wood | China | KY494714 | KY806200 | KY705113 | |
A. xenocordella |
|
Soil from roadway | Zimbabwe | KF144925 | KF145013 | KF145055 |
LC3486 | Camellia sinensis | China | KY494687 | KY705158 | KY705086 | |
A. yunnanum |
|
Decaying bamboo culms | China | KU940147 | – | – |
N. gorlenkoana |
|
Vitis vinifera | Kazakhstan | KX986048 | KY019456 | KY019420 |
To determine the distribution of Arthrinium species on host/substrate, the number of species occurred on each host (based on family level) was counted based on data from this study, relevant literature and the USDA fungal database (https://nt.ars-grin.gov/fungaldatabases/). The proportion account for the known 66 species in Arthrinium (Index Fungorum) was illustrated in a histogram. Four species with an unknown host range were not included in this analysis.
The combined ITS, TUB2 and TEF1 dataset contained 75 strains, with Nigrospora gorlenkoana
The histogram in Figure
CHINA, Guangdong Province, on bamboo leaves, 10 Jul. 2016, D.W. Xiao, (holotype:
Named after the host of the holotype.
Hyphae hyaline, branched, septate, 1.5–5.0 μm diam. Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect, aggregated in clusters on hyphae, hyaline to pale brown, smooth, doliiform to ampulliform, or lageniform, 4.0–12.0 × 3.0–7.0 μm (x̄ = 6.6 ± 1.8 × 4.8 ± 0.9, n = 30). Conidia olivaceous to brown, smooth to finely roughened, subglobose to ellipsoid, 11.5–15.5 × 7.0–14.0 μm (x̄ = 13.2 ± 0.8 × 11.4 ± 1.2, n = 50).
On PDA, colonies flat, spreading, margin circular, with abundant aerial mycelia, surface and reverse white to grey. On MEA, colonies flat, spreading, surface and reverse brown to black.
CHINA, Jiangxi Province, on bamboo leaves, 10 Jul. 2016, Q. Xiong, living culture LC7246; Guangdong Province, on bamboo leaves, 10 Jul. 2016, D.W. Xiao, living culture LC7107; ibid. living culture LC7113; ibid. living culture LC7124; ibid. living culture LC7125; ibid. living culture LC7128.
Seven strains representing A. bambusae clustered in a well-supported clade closely related to A. subroseum (98% sequence similarity in ITS; 92% in TUB2; 96% in TEF1). Arthrinium bambusae differs from A. subroseum in the morphology of conidiophore (reduced to conidiogenous cells in A. bambusae vs. erect or ascending, clustered in groups in A. subroseum). Moreover, A. bambusae does not produce pigment on the PDA.
CHINA, Jiangxi Province, on Camellia sinensis, 22 Apr. 2013, Q. Chen, (holotype:
Named with the host plant of the type.
Hyphae hyaline, branched, septate, 2.0–4.5 μm diam. Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect, aggregated in clusters, hyaline to pale brown, smooth, doliiform to ampulliform, 4.0–9.5 × 3.0–6.0 μm (x̄ = 6.1 ± 1.4 × 4.4 ± 0.9, n = 30). Conidia brown to dark brown, smooth, globose to subglobose, 9.0–13.5 × 7.0–12.0 μm (x̄ = 11.1 ± 0.9 × 10.1 ± 1.0, n = 50).
On PDA, colonies flat, margin circular, initially white, becoming greyish on surface, reaching 9 cm in 7 days at 25 °C. On MEA, with sparse aerial mycelia, surface dirty white, reverse pale luteous.
CHINA, Hubei Province, on Brassica campestris, 31 Mar. 2016, Y.Z. Zhao, living culture LC8181 = LF1498.
Two strains representing A. camelliae-sinensis clustered in a well-supported clade and appeared closely related to A. jiangxiense (97% sequence similarity in ITS; 94% in TUB2; 94% in TEF1) and A. obovatum (98% sequence similarity in ITS; 95% in TUB2; 93% in TEF1). While A. camelliae-sinensis is distinct from A. jiangxiense in its larger conidia (globose or subglobose, 9.0–13.5 × 7.0–12.0 μm in A. camelliae-sinensis vs. surface view 7.5–10.0 μm diam, side view 4.5–7.0 μm diam in A. jiangxiense) and conidiogenous cell arrangement (aggregated irregularly on hyphae vs. scattered on hyphae in A. jiangxiense) and distinct from A. obovatum in the lack of obovoid conidia (see the note under A. obovatum).
CHINA, Chongqing, on Dichotomanthus tristaniaecarpa, 20 Dec. 2012, L. Cai, (holotype:
Named after the host from which it was isolated.
Hyphae hyaline, branched, septate, 1.5–5.0 μm diam. Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect, aggregated in clusters on hyphae, hyaline to pale brown, smooth, doliiform to clavate or lageniform, 5.5–11.0 × 3.0–5.0 μm (x̄ = 7.9 ± 1.4 × 4.0 ± 0.5, n = 30). Conidia brown to dark brown, smooth to finely roughened, globose, subglobose to lenticular, with a longitudinal germ slit, 9.0–15.0 × 6.0–12.0 μm (x̄ = 12.0 ± 1.4 × 8.5 ± 1.1, n = 50).
On PDA, colonies umbonate, margin irregular, with sparse aerial mycelia. Colonies creamy-white to greyish without patches reverse, reaching 9 cm in 7 days at 25 °C. On MEA, colonies flat, spreading, surface and reverse pale luteous.
CHINA, Chongqing, on Dichotomanthus tristaniaecarpa, 20 Dec. 2012, L. Cai, living culture LC8175 = WM529; ibid. living culture LC8176 = WM 530.
Three strains representing A. dichotomanthi formed a distinct clade closely related to A. phaeospermum (Corda) M.B. Ellis (99% sequence similarity in ITS; 96% in TUB2; 96% in TEF1), A. serenense Larrondo & Calvo (99% sequence similarity in ITS; 95% in TUB2) and A. saccharicola F. Stevens (99% sequence similarity in ITS; 95% in TUB2; 97% in TEF1). Arthrinium dichotomanthi differs from A. phaeospermum and A. saccharicola in its larger conidia (globose or subglobose, 9.0–15.0 × 6.0–12.0 μm in A. dichotomanthi vs. surface view (9–)10(–12) μm diam, side view 6–7 μm diam in A. phaeospermum, surface view (7–)8–9(–10) μm diam, side view (4–)5(–6) μm diam in A. saccharicola) and from A. serenense by the absence of odour on the MEA colony (Larrondo 1990).
CHINA, Guizhou Province, from the air in karst cave, 23 Jul. 2014, Z.F. Zhang, (holotype:
Named after the province where type was collected, Guizhou province.
Hyphae hyaline, branched, septate, 1.5–5.0 μm diam. Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect, aggregated in clusters on hyphae, pale brown, smooth, subglobose, ampulliform or doliiform, 3.5–8.0 × 3.0 – 4.5 μm (x̄ =5.1 ± 1.08 × 3.7 ± 0.49, n = 30). Conidia dark brown to black, smooth to finely roughened, globose or subglobose, occasionally elongated to ellipsoidal, with a longitudinal, hyaline, thin, germ slit, 5.0–7.5 × 4.0–7.0 μm (x̄ = 6.1 ± 0.5 × 5.5 ± 0.6, n = 50).
On PDA, colonies flat, woolly, margin circular, with moderate aerial mycelia, surface initially white, becoming greyish and reverse with black patches, reaching 9 cm in 9 days at 25 °C. On MEA, surface dirty white with patches of olivaceous-grey and reverse greyish.
CHINA, Guizhou Province, from the air in karst cave, 23 Jul. 2014, Z.F. Zhang, living culture LC5318.
Arthrinium guizhouense is closely related to A. sacchari (Speg.) M.B. Ellis (99% sequence similarity in ITS; 99% in TUB2; 94% in TEF1). Morphologically, A. guizhouense and A. sacchari are very similar in conidial size, but A. guizhouensis produces relatively shorter conidiogenous cells (3.5–8.0 μm in A. guizhouense vs. 5–12 μm in A. sacchari).
CHINA, Jiangxi Province, on Maesa sp., 05 Sept. 2013, Y.H. Gao, (holotype:
Named after the province where the most strains of this species were collected, Jiangxi.
Hyphae hyaline, branched, septate, 1.5–5.0 μm diam. Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect, scattered or aggregated in clusters on hyphae, hyaline to pale brown, smooth, ampulliform, 6.0–15.0 × 2.5–5.0 μm (x̄ = 9.7 ± 2.6 × 3.7 ± 0.6, n = 30), apical neck 2.5–6.0 μm long, basal part 3.0–9.0 μm long. Conidia brown, smooth to finely roughened, granular, globose to ellipsoid in surface view, 7.5–10.0 μm diam (x̄ = 8.7 ± 0.6, n = 50), lenticular in side view, with longitudinal, pale germ slit, 4.5–7.0 μm diam (x̄ = 5.8 ± 0.6, n = 50). Sterile cells forming on solitary loci on hyphae, brown, finely roughened, subcylindrical to clavate.
On PDA, colonies flat, woolly, margin circular, with sparse aerial mycelia, initially white, becoming greyish due to sporulation, reaching 9 cm in 10 days at 25 °C, on MEA, sienna with patches of luteous, reverse luteous to sienna.
CHINA, Hunan Province, on bamboo, 22 Sept. 2010, L. Cai, living culture LC2831; Jiangxi Province, on Phyllostachys sp., 05 Sept. 2013, Y.H. Gao, living culture LC4494; on Phyllostachys sp., 22 Apr. 2013, Q. Chen, living culture LC4993; ibid. living culture LC4497; ibid. living culture LC5001; ibid. living culture LC5004; on Imperata cylindrical, 22 Apr. 2013, Q. Chen, living culture LC5015; on Maesa sp., 05 Sept. 2013, Y.H. Gao, living culture LC4541; on Machilus sp., 05 Sept. 2013, Y.H. Gao, living culture LC4547; on Camellia sinensis, 05 Sept. 2013, Y.H. Gao, living culture LC4578; on bamboo, 01 Jul. 2016, J.E. Huang, living culture LC7104; ibid. living culture LC7154; ibid. living culture LC7156; ibid. living culture LC7275.
Two strains representing Arthrinium jiangxiense clustered in a well-supported clade and appeared closely related to A. camelliae-sinensis (97% sequence similarity in ITS; 94% in TUB2; 94% in TEF1). While A. jiangxiensis is distinct from A. camelliae-sinensis in its smaller conidia (surface view 7.5–10.0 μm diam, side view 4.5–7.0 μm diam in A. jiangxiensis vs. globose or subglobose, 9.0–13.5 × 7.0–12.0 μm in A. camelliae-sinensis) and conidiogenous cell arrangements (conidiogenous cells scattered on hyphae vs. aggregated irregularly on hyphae in A. jiangxiense).
CHINA, Chongqing, on Lithocarpus sp., 20 Dec. 2012, L. Cai, (holotype:
Referring to the production of the large obovoid conidia.
Hyphae hyaline to pale brown, branched, septate, 1.5–5.0 μm diam. Conidiophores reduced to conidiogenous cells. Conidiogenous cells erect, aggregated in clusters on hyphae, pale brown, smooth, subcylindrical or clavate, 5.5–13.5 × 2.5–5.0 μm (x̄ = 8.7 ± 2.4 × 3.6 ± 0.6, n = 30). Conidia dark brown, roughened, globose to subglobose, 11.0–16.5 μm (x̄ = 13.8 ± 1.5, n = 50) in diam.; obovoid, 16.0–31.0 × 9.0–16.0 μm (x̄ = 23.0 ± 2.7 × 12.7 ± 1.4, n = 50), occasionally elongated to ellipsoidal.
On PDA, colonies flat, spreading, margin circular, initially white, becoming olivaceous-grey on surface, reverse smoke-grey with patches of olivaceous grey, reaching 9 cm in 7 days at 25 °C. On MEA, surface olivaceous grey in the central and luteous around, reverse with patches of olivaceous grey.
CHINA, Chongqing, on Lithocarpus sp., 20 Dec. 2012, L. Cai, living culture LC8177; ibid. living culture LC8178.
Arthrinium obovatum is the only species that produces obovoid conidia (Figure
CHINA, Guangdong Province, on bamboo, Jul. 2016, D.W. Xiao, (holotype:
Referring to the pseudoparenchymatous hyphae.
Hyphae hyaline to pale brown, branched, septate, 1.5–5.0 μm diam., pseudoparenchymatous. Conidiophores aggregated in hyaline to light brown sporodochia, smooth, usually unbranched, up to 40 μm long, 3–6 μm width. Conidiogenous cells hyaline to pale yellow, smooth to finely roughened, subcylindrical to doliiform, 8.0–18.5 × 3.0–8.5μm (x̄ = 13.7 ± 3.2 × 5.4 ± 1.2, n = 30). Conidia pale to dark brown, smooth, finely guttulate, globose to subglobose, 13.5–27.0 × 12.0–23.5 μm (x̄ = 20.2 ± 2.5 × 17.1 ± 2.4, n = 50). Sometimes lobed or dentate, polygonal or irregular in surface view.
On PDA, colonies flat, spreading, margin circular, with moderate aerial mycelia, initially white, becoming grey on surface, reverse smoke-grey without patches, reaching 9 cm in 8 days at 25 °C. On MEA, surface pale luteous to grey with abundant mycelia, reverse greyish without patches.
CHINA, Guangdong Province, on bamboo, Jul. 2016, D.W. Xiao, living culture LC8173; ibid. living culture LC8174.
Arthrinium pseudoparenchymaticum is closely related to A. hyphopodii (94% sequence similarity in ITS), but differs in its much larger conidia (13.5–27.0 × 12.0–23.5 μm vs. 5–10 × 4–8 μm), the absence of hyphopodia and the presence of dentate conidia.
CHINA, Jiangxi Province, on bamboo, 1 Jul. 2016, J.E. Huang, (holotype:
Named after the colour of colony on PDA, pinkish.
Hyphae hyaline to pale brown, branched, septate, 1.5–6.0 μm diam. Conidiophores hyaline to pale brown, smooth, erect or ascending, simple, flexuous, subcylindrical, clustered in groups. Conidiophores aggregated in brown sporodochia, smooth, hyaline to brown, up to 20 μm long, 2–4.5 μm width. Conidiogenous cells pale brown, smooth, doliiform to subcylindrical, 3.0–6.5 × 2.0–5.0 μm (x̄ = 4.7 ± 1.2 × 3.7 ± 0.9, n = 30). Conidia pale brown to dark brown, smooth, globose to subglobose or ellipsoidal, 12.0–17.5 × 9.0–16.0 μm (x̄ = 14.9 ± 1.4 × 11.8 ± 1.8, n = 50).
On PDA, colonies flat, spreading, margin circular, with moderate aerial mycelia, initially white, becoming light pink on surface, reverse peach-puff without patches, reaching 10 cm in 8 days at 25 °C. On MEA, surface blackish-green with abundant mycelia, reverse with patches of greyish.
CHINA, Jiangxi Province, on bamboo, 1 Jul. 2016, J.E. Huang, living culture LC7215; ibid. living culture LC7291.
Three strains representing A. subroseum clustered in a well-supported clade, closely related to A. garethjonesii (94% sequence similarity in ITS) and A. bambusae (98% sequence similarity in ITS; 92% in TUB2; 96% in TEF1). However, A. subroseum differs from A. bambusae in the morphology of conidiophores (erect or ascending, clustered in groups in A. subroseum vs. reduced to conidiogenous cells in A. bambusae). Arthrinium subroseum is not morphologically comparable to A. garethjonesii, whose asexual morph is undetermined (
Arthrinium, Cordella and Pteroconium share similar morphological characters, e.g. basauxic conidiophores with terminal and intercalary polyblastic conidiogenous cells and brown, unicellular conidia with a pallid germ slit (
Currently there are 70 recognised species in Arthrinium (Index Fungorum), occurring on a wide variety of both living and decaying plant materials. It is noteworthy that Arthrinium species showed distinct preference for growing on two graminaceous families, Poaceaeand Cyperaceae, amongst which, Bambusa (Poaceae) and Carex (Cyperaceae) are two of the most common host genera for Arthrinium species. For example, seven species have been recorded from Carex spp., i.e. A. austriacum Petr. (1959), A. caricicola Kunze (1817), A. globosum
In addition to the Arthrinium species from China, we also tried to resolve the phylogenetic status of Arthrinium mytilomorphum Bhat & W.B. Kendr. (
Teleomorph-typified genus Apiospora was treated as a synonym of anamorph-typified genus Arthrinium on the basis that Arthrinium is older and more commonly used in literature (Crous et al. 2013). However, only three of the 58 recorded Apiospora species have been properly linked to their known Arthrinium counterparts, i.e. Arthrinium hysterinum (syn. Ap. bambusae) (
We thank Peng Zhao, Qian Chen, Yahui Gao and Zhifeng Zhang for providing strains and technical assistance. We kindly appreciated the curator of Agriculture and Agri-Food Canada herbarium and Dr. Wen Chen in Ottawa Research and Development Centre AAFC for providing DNA samples and microscope slides of the type specimen of Arthrinium mytilomorphum. This work was financially supported by the National Science Fund for Distinguished Young Scholars of China (NSFC 31725001) and the Frontier Science Research Project of the Chinese Academy of Sciences (QYZDB-SSW-SMC044).