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e genera Rugonectria and elonectria in China 101
The genera Rugonectria and Thelonectria
(Hypocreales, Nectriaceae) in China
Zhao-Qing Zeng1, Wen-Ying Zhuang1
1 State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China
Corresponding authors: Zhao-Qing Zeng (zengzq@im.ac.cn); Wen-Ying Zhuang (zhuangwy@im.ac.cn)
Academic editor: Danny Haelewaters|Received14 March 2019|Accepted 22 May 2019|Published 1 July2019
Citation: Zeng Z-Q, Zhuang W-Y (2019) e genera Rugonectria and elonectria (Hypocreales,Nectriaceae) in
China. MycoKeys 55: 101–120. https://doi.org/10.3897/mycokeys.55.34527
Abstract
Recent collections and herbarium specimens of Rugonectria and elonectria from dierent regions of China
were examined. Using combined analyses of morphological and molecular data, 17 species are recognised
including three species of Rugonectria and 14 species in elonectria. Amongst them, R. microconidia
and T. guangdongensis are new to science. Rugonectria microconidia on mossy bark is characterised by
supercial, yellow to orange, pyriform to subglobose perithecia with a warted surface; ellipsoidal to
broadly ellipsoidal, striate, uniseptate ascospores; and allantoid to rod-shaped, aseptate microconidia.
elonectria guangdongensis possesses bright red perithecia with a slightly roughened surface and a
prominently dark papilla; ellipsoidal, smooth, uniseptate ascospores; and subcylindrical, slightly curved,
multiseptate macroconidia. Morphological distinctions and sequence divergences between the new species
and their close relatives are discussed. Name changes for the previously recorded species in China are noted.
Keywords
Morphology, Multigene analyses, Taxonomy
Introduction
e family Nectriaceae was introduced in 1865 and circumscribed to accommodate
the hypocrealean species having ascomata that are generally yellow, orange-red to pur-
ple and usually changing colour in potassium hydroxide (KOH) and lactic acid (LA)
(Rossman et al. 1999). About 55 genera containing 900 species are included in the
family (Lombard et al. 2015). A phylogenetic backbone for Nectriaceae was construct-
ed based on DNA sequences of 10 loci by Lombard et al. (2015).
Copyright Zhao-Qing Zeng, Wen-Ying Zhuang. This is an open access article distributed under the terms of the Creative Commons Attribution License
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MycoKeys 55: 101–120 (2019)
doi: 10.3897/mycokeys.55.34527
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RESEARCH ARTICLE
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
102
e genus Rugonectria P. Chaverri & Samuels, typied by R. rugulosa (Pat. & Gaillard)
Samuels, P. Chaverri & C. Salgado, is characterised by perithecia solitary or in groups,
seated on or partially immersed in a stroma. e perithecia are orange to red, globose to
subglobose and non-papillate, with warted or rugose walls. Ascospores are ellipsoidal to
oblong, striate, hyaline and 1-septate; and microconidia are ovoid to cylindrical (Chaverri
et al. 2011). Currently, four species are recognised in the genus (Chaverri et al. 2011;
Zeng et al. 2012). elonectria P. Chaverri & C. Salgado, typied by T. discophora (Mont.)
P. Chaverri & C. Salgado, was established by Chaverri et al. (2011) to accommodate the
nectriaceous fungi having supercial, globose to subglobose or pyriform to elongated
perithecia which do not collapse when dry, with a prominent and darkened papilla;
smooth, rarely spinulose or striate ascospores and curved macroconidia with rounded ends
(Chaverri et al. 2011; Lombard et al. 2015; Salgado-Salazar et al. 2016). About 44 species
are currently accepted in the genus (Chaverri et al. 2011; Salgado-Salazar et al. 2012,
2015, 2016; Zeng and Zhuang 2013; Crous et al. 2018). Species in the genera Rugonectria
and elonectria are distributed in the tropics, subtropics and temperate regions and occur
on early decaying bark, roots, branches, trunks and rarely in soil (Chaverri et al. 2011;
Salgado-Salazar et al. 2015). A few species are plant pathogenic, such as R. castaneicola
(W. Yamam. & Oyasu) Hirooka & P. Chaverri causing Abies and Acer cankers and T.rubi
(Osterw.) C. Salgado & P. Chaverri causing Rubus cankers (Cedeño et al. 2004; Kobayashi
et al. 2005; Chaverri et al. 2011; Salgado-Salazar et al. 2015).
e rst record of Rugonectria from China dates back to 2000 when R. rugulosa (as
Nectria rugulosa Pat. & Gaillard) was reported by Lu et al. (2000) based on a specimen
collected on dead petioles of king palm. Research on elonectria in China was started
by Teng (1936) when T. discophora (as N. discophora Mont.) was rst reported on bark
of fallen branches from Yunnan Province. In connection with our current work on the
Chinese fungus ora, fresh materials and herbarium specimens of the two genera were
examined. Based on morphology and phylogenetic analyses of the partial sequences
of α-actin (ACT), internal transcribed spacer (ITS), nuclear ribosomal large subunit
(LSU) rDNA and the largest subunit of RNA polymerase II (RPB1), 17 species were
identied, including two undescribed species. Morphological and molecular diagnos-
tic features between the new taxa and their closely related fungi are discussed.
Materials and methods
Sampling and morphological studies
Specimens were collected from Beijing, Fujian, Guangdong, Hainan, Henan, Hubei,
Hunan and Yunnan provinces and are deposited in Herbarium Mycologicum Academiae
Sinicae (HMAS) and cultures are kept in the State Key Laboratory of Mycology, Institute
of Microbiology, Chinese Academy of Sciences. e methods used by Luo and Zhuang
(2010) and Chaverri et al. (2011) were followed for morphological observations. e
ascomatal wall reactions to 3% KOH and 100% LA were tested. To observe micro-
e genera Rugonectria and elonectria in China 103
morphological characteristics of perithecial walls, sections were made with a freezing
microtome (YD-1508-III, Jinhua, China) at a thickness of 6–8 μm. Lactophenol cotton
blue solution was used as mounting medium for examination of anatomic structures and
measurements of perithecia, asci and ascospores. Photographs were taken with a Leica
DFC450 digital camera (Wetzlar, Germany) attached to a Leica M125 stereomicroscope
(Milton Keynes, UK) for gross morphology and a Zeiss AxioCam MRc 5 digital cam-
era (Jena, Germany) attached to a Zeiss Axio Imager A2 microscope (Göttingen, Ger-
many) for microscopic features. Descriptive statistics of ascospores and conidia (mini-
mum, maximum, mean and standard deviation) were calculated following the methods
of Hirooka et al. (2012). Measurements of individual structures were based on 30 units,
except as otherwise noted. Morphology of colonies were characterised using potato dex-
trose agar (PDA, 20% w/v potato + 2% w/v dextrose + 2% w/v agar) and synthetic
nutrient-poor agar (SNA; Nirenberg 1976) at 25 °C in an incubator with alternating
periods of light and darkness (12 h/12 h). Colony growth rates were measured after 7 d.
DNA extraction, PCR amplification and sequencing
Genomic DNA was extracted from fresh mycelium following the method of Wang and
Zhuang (2004). Four primer pairs, act1-act2 (Samuels et al. 2006), ITS5-ITS4 (White et
al. 1990), LR0R-LR5 (Vilgalys and Hester 1990; Rehner and Samuels 1994) and crpb1a-
rpb1c (Castlebury et al. 2004) were used to amplify the ACT, ITS, LSU and RPB1 regions,
respectively. PCR reactions were performed using an ABI 2720 ermal Cycler (Applied
Biosciences, Foster City, USA) with a 25 μl reaction system consisting of 12.5 μl Taq Mas-
terMix, 1 μl each primer (10 μM), 1 μl template DNA and 9.5 μl ddH2O, based on the
procedures detailed in Chaverri et al. (2011). DNA sequencing was carried out in both
directions on an ABI 3730XL DNA Sequencer (Applied Biosciences, Foster City, USA).
Sequence alignment and phylogenetic analyses
Newly obtained sequences and those retrieved from GenBank are listed in Table I.
e sequences were assembled, aligned and the primer sequences were trimmed via
BioEdit 7.0.5 (Hall 1999) and converted to NEXUS les by ClustalX 1.8 (ompson
et al. 1997). A partition homogeneity test was performed with 1,000 replicates in
PAUP*4.0b10 (Swoord 2002) to evaluate statistical congruence amongst the four
loci. e aligned ACT, ITS, LSU and RPB1 sequences were combined in BioEdit and
analysed with Bayesian Inference (BI), Maximum Parsimony (MP) and Maximum
Likelihood (ML) methods to determine the phylogenetic positions of the new species.
e MP analysis was performed with PAUP 4.0b10 (Swoord 2002) using 1000 repli-
cates of heuristic search with random addition of sequences and subsequent TBR (tree
bisection and reconnection) branch swapping. Topological condence of the resulting
trees was tested by Maximum Parsimony bootstrap proportion (MPBP) with 1000
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
104
Table 1. List of species, herbarium/strain numbers and GenBank accession numbers of materials
used in this study.
Species Herbarium/strain
numbers
GenBank Accession numbers
ACT ITS LSU RPB1
Cosmospora coccinea Rabenh. CBS 114050 GQ505967 FJ474072 GQ505990 GQ506020
Nectria cinnabarina (Tode) Fr. AR 4302/AR 4477 HM484627 HM484548 HM484562 HM484577
Rugonectria castaneicola (W. Yamam. & Oyasu) Hirooka &
P. Chaverri
CBS 128360 –MH864901 MH876352 –
R. microconidia Z.Q. Zeng & W.Y. Zhuang HMAS 254521 MF669044 a MF669050 MF669052 MF669056
R. neobalansae (Samuels) P. Chaverri & Samuels CBS 125120 –KM231750 HM364322 KM232146
R. rugulosa (Pat. & Gaillard) Samuels, P. Chaverri & C.
Salgado
YH 1001 JF832515 JF832661 JF832761 JF832836
R. sinica W.Y. Zhuang, Z.Q. Zeng & W.H. Ho HMAS 183542 MF669046 HM054141 HM042430 MF669058
elonectria asiatica C. Salgado & Hirooka MAFF 241576 KC121436 KC153774 KC121500 KC153967
T. beijingensis Z.Q. Zeng, J. Luo & W.Y. Zhuang HMAS 188498 MF669047 JQ836656 MF669054 MF669059
T. blattea C. Salgado & P. Chaverri CBS 95268 KC121387 KC153725 KC121451 KC153918
T. brayfordii C. Salgado & Samuels CBS 118612 KC121381 KC153719 KC121445 KC153912
T. conchyliata C. Salgado & P. Chaverri GJS 8745 KC121401 KC153739 KC121465 KC153932
T. discophora (Mont.) P. Chaverri & C. Salgado AR 4742 KC121376 KC153714 KC121440 KC153907
T. guangdongensis Z.Q. Zeng & W.Y. Zhuang HMAS 254522 MF669045 MF669051 MF669053 MF669057
T. ianthina C. Salgado & Guu GJS 10118 KC121393 KC153731 KC121457 KC153924
T. japonica C. Salgado & Hirooka MAFF 241524 KC121428 KC153766 KC121492 KC153959
HMAS 98327 MK556799 HM054140 HM042434 –
T. mammoidea (W. Phillips & Plowr.) C. Salgado & R.M.
Sanchez
IMI 69361 KC121425 KC153763 KC121489 KC153956
T. ostrina C. Salgado & P. Chaverri GJS 9623 KC121418 KC153756 KC121482 KC153949
T. phoenicea C. Salgado & P. Chaverri GJS 85179 KC121398 KC153736 KC121462 KC153929
HMAS 76856 MK556800 JQ836657 DQ119572 –
T. pinea (Dingley) C. Salgado & P. Chaverri AR 4324 HM352875 HM364294 HM364307 HM364326
T. porphyria C. Salgado & Hirooka MAFF 241515 KC121426 KC153764 KC121490 KC153957
HMAS 98333 MK556798 HM054136 HM042433 –
T. purpurea C. Salgado & P. Chaverri GJS 10131 KC121394 KC153732 KC121458 KC153925
T. rubi (Osterw.) C. Salgado & P. Chaverri CBS 11312 KC121380 KC153718 KC121444 KC153911
T. sinensis (J. Luo & W.Y. Zhuang) Z.Q. Zeng & W.Y.
Zhuang
HMAS 183186 MF669048 FJ560441 FJ560436 MF669060
T. tyrus C. Salgado & P. Chaverri GJS 9046 KC121413 KC153751 KC121477 KC153944
T. violaria C. Salgado & R.M. Sanchez AR 4766 KC121377 KC153715 KC121441 KC153908
T. yunnanica Z.Q. Zeng & W.Y. Zhuang HMAS 183564 MF669049 FJ560438 MF669055 MF669061
a e GenBank numbers in bold type were newly generated in this study.
replications, each with 10 replicates of random addition of taxa. e BI analysis was
conducted by MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003) using a Markov chain
Monte Carlo algorithm. Nucleotide substi tution models were determined by MrMod-
eltest 2.3 (Nylander 2004). Four Markov chains were run simultaneously for 1000000
generations with the trees sam pled every 100 generations. A 50% majority rule con-
sensus tree was computed after excluding the rst 2500 trees as ‘burn-in’. Bayesian
Inference posterior probability (BIPP) was determined from the remaining trees. ML
analysis was conducted with IQ-Tree 1.6.10 (Nguyen et al. 2015) using the best model
for each locus chose by ModelFinder (Chernomor et al. 2016). Branch support meas-
ures were calculated with 1000 bootstrap replicates. Trees were examined by TreeView
1.6.6 (Page 1996). Cosmospora coccinea Rabenh. and Nectria cinnabarina (Tode) Fr.
were used as outgroup taxa. Maximum Likelihood bootstrap proportion (MLBP) and
MPBP greater than 50% and BIPP greater than 90% were shown at the nodes.
e genera Rugonectria and elonectria in China 105
Results
e sequences of ACT, ITS, LSU and RPB1 from 25 representative taxa of Rugonectria
and elonectria were analysed. e partition homogeneity test (P = 0.03) indicated
that the individual partitions were not highly incongruent (Cunningham 1997), thus
these four loci were combined for the phylogenetic analyses. In the MP analysis, the
datasets included 2524 nucleotide characters, of which 1836 were constant, 198 were
variable and parsimony-uninformative and 490 were parsimony-informative. e MP
analysis resulted in three most parsimonious trees (tree length = 1415, CI = 0.6721,
HI = 0.3279, RI = 0.6098, RCI = 0.5351). One of them is shown in Figure 1. e
ML and BI trees were of similar topology. e nal matrix was deposited in Tree-
BASE with accession no. S23994. e isolate HMAS 254521 grouped with other
members of Rugonectria by receiving high bootstrap values (MLBP/MPBP/BIPP =
100%/100%/100%) and the isolate HMAS 254522 clustered with the representatives
of elonectria (MLBP/MPBP/BIPP = 100%/100%/100%), which support the taxo-
nomic placements of these new species.
Taxonomy
Rugonectria microconidia Z.Q. Zeng & W.Y. Zhuang, sp. nov.
Fungal Names: FN570487
Figure 2
Holotype. CHINA. Hunan, Yizhang, Mangshan, (24°57'56.58"N, 112°57'34.63"E),
alt. 700 m, on mossy bark, 26 October 2015, Z.Q. Zeng, X.C. Wang, K. Chen, Y.B.
Zhang 10266 (HMAS 254521); ex-type culture: HMAS 247232.
Sequences. ACT (MF669044), ITS (MF669050), LSU (MF669052) and RPB1
(MF669056).
Etymology. e specic epithet refers to the microconidia produced in culture.
Description. Mycelium not visible around ascomata or on natural substrata. Asco-
mata supercial, gregarious, with basal stroma, pyriform to subglobose, non-papillate,
yellow to orange, often with a darker red ostiolar area when dry, turning dark red in
KOH, becoming slightly yellow in LA, 421–549 × 333–470 μm (n = 8). Perithecial
surface warted, 30–93 μm thick, of textura globulosa to textura angularis, cells 10–27×
8–18 μm, walls 1.5–2.5 μm thick. Perithecial wall of two layers, 45–70 μm thick, outer
layer 25–45 μm thick, of textura globulosa to textura angularis; inner layer 7–25 μm
thick, of textura prismatica. Asci unitunicate, clavate, 8-spored, 93–130 × (11–)15–25
μm (112.6 ± 12.6 × 18.9 ± 3.2 μm). Ascospores ellipsoid to broadly ellipsoid, 1-sep-
tate, striate, uniseriate or biseriate above and uniseriate below, hyaline, 20–28 × 8–12
μm (24.0 ± 2.0 × 10.1 ± 0.9 μm). Colony on PDA 42 mm diameter after 7 d under
daylight at 25 °C, surface velvety, with white aerial mycelium, producing pale pinkish
pigment in medium. Colony on SNA reaches 40 mm diameter after 7 d under daylight
at 25 °C, surface with sparse whitish aerial mycelium. Conidiophores simply branched,
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
106
Figure 1. A Maximum Parsimony tree inferred from the combined ACT, ITS, LSU and RPB1 sequences.
Cosmospora coccinea and Nectria cinnabarina were used as outgroup taxa. MLBP (left) and MPBP (mid-
dle) above 50%, BIPP (right) above 90% are indicated at nodes.
18–50 × 2–3 μm. Microconidia allantoid to rod shaped, slightly curved, 0(1–2)-sep-
tate, 3–14(–18) × 1.2–2.5(–3) μm (6.7 ± 3.1 × 1.6 ± 0.4 μm).
Habitat. On mossy bark.
Distribution. Asia (China).
e genera Rugonectria and elonectria in China 107
Figure 2. Rugonectria microconidia a–d ascomata on natural substratum e colony on PDA f colony on
SNA g, h median section through perithecium i–k asci with ascospores l–o ascospores p–s conidiophores
and conidia t, u conidiogenous cells and conidia v, w microconidia. Scale bars: 0.5 mm (a–d); 50 μm
(g, h); 10 μm (i–w).
Notes. e non-papillate perithecia with warted surface, clavate asci with ellipsoi-
dal to broadly ellipsoidal, uniseptate, striate ascospores, as well as our molecular data,
suggest that this species belongs to Rugonectria (Chaverri et al. 2011). Amongst the
known species of the genus, R. microconidia is morphologically most similar to the type
species, R. rugulosa, in having gregarious, warted, orange perithecia often with a dark
red ostiole when dry (Samuels et al. 1990; Samuels and Brayford 1994). e newly de-
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
108
scribed species diers in having asci that are 93–130 × (11–)15–25 μm and larger than
those of R. rugulosa that are (53–)64–83(–95) × (7.5–)11.3–15.5(–17) μm. In addi-
tion, the ascospores of R. microconidia are also larger, 20–28 × 8–12 μm, while those of
R.rugulosa are (10–)13.5–18(–24) × (3.3–)4.7–6.7(–10) μm. Unlike R. microconidia,
R. rugulosa does not produce macroconidia in culture (Samuels et al. 1990; Samuels
and Brayford 1994). Sequence comparisons reveal that there are 21 bp, 21 bp, 12 bp
and 22 bp divergences in the ACT, ITS, LSU and RPB1 regions, respectively, between
R. microconidia and R. rugulosa (YH1001). Both morphological and molecular data
suggest that these species are distinct.
Rugonectria rugulosa (Pat. & Gaillard) Samuels, P. Chaverri & C. Salgado, in
Chaverri, Salgado, Hirooka, Rossman & Samuels, Stud. Mycol. 68: 73, 2011
≡ Nectria rugulosa Pat. & Gaillard, Bull. Soc. Mycol. Fr. 5(4): 115, 1890.
≡ Neonectria rugulosa (Pat. & Gaillard) Mantiri & Samuels, in Mantiri, Samuels, Rahe
& Honda, Can. J. Bot. 79(3): 339, 2001.
= Cylindrocarpon rugulosum Brayford & Samuels, in Samuels & Brayford, Sydowia
46(1): 148, 1994.
Specimens examined. CHINA. Henan, Jigongshan, alt. 400 m, on rotten twigs, 14
November 2003, W.Y. Zhuang, Y. Nong 5142 (HMAS 91774). Hainan, Changjiang,
Bawangling, alt. 1100 m, on rotten twigs, 7 December 2000, W.Y. Zhuang, X.M.
Zhang H25 (HMAS 83349); Ledong, Jianfengling, alt. 1100 m, on rotten twigs, 9
December 2000, W.Y. Zhuang, X.M. Zhang, Z.H. Yu H36, H41 (HMAS 83350,
83370); Qiongzhong, Limushan, alt. 700 m, on rotten twigs, 18 December 2000,
W.Y. Zhuang, X.M. Zhang H124 (HMAS 76867); Tongzha, Wuzhishan, alt. 1000 m,
on bark, 16 December 2000, W.Y. Zhuang, X.M. Zhang, Z.H. Yu, Y.H. Zhang H105
(HMAS 83371); on rotten twigs, W.P. Wu W7058 (HMAS 183161); Yunnan, Xichou,
on rotten twigs, 11 November 1999, W.Y. Zhuang, Z.H. Yu 3407 (HMAS 183160).
Habitat. On rotten twigs, wood of recently dead and dying trees.
Distribution. Africa (Congo), Americas (Venezuela), Asia (China, Indonesia),
possibly pantropical.
Notes. e species was formerly placed in Nectria (Fr.) Fr. and Neonectria Wol-
lenw. until Chaverri et al. (2011) introduced Rugonectria with R. rugosa as the type
species. e Chinese materials match well the description of the fungus (Samuels and
Brayford 1994).
Rugonectria sinica W.Y. Zhuang, Z.Q. Zeng & W.H. Ho, in Zeng, Zhuang & Ho,
Mycosystema 31(4): 467, 2013
Specimens examined. CHINA. Hainan, Changjiang, Bawanling, alt. 1100 m, on
dead twigs of Quercus sp., 7 December 2000, W.Y. Zhuang, X.M. Zhang H22, H30
e genera Rugonectria and elonectria in China 109
(HMAS 76854, 83369); Changjiang, Bawanling, alt. 1100 m, on dead twigs, 7 De-
cember 2000, W.Y. Zhuang, X.M. Zhang H28 (HMAS 76865); Lingshui, Diaolu-
oshan, alt. 1100 m, on bark, 13 December 2000, W.Y. Zhuang, X.M. Zhang, Z.H. Yu
H70 (HMAS 76866); Henan, Jigongshan, alt. 400 m, on dead twigs, 14 November
2003, W.Y. Zhuang, Y. Nong 5099 (HMAS 91773); Fujian, Wuyishan, on dead twigs,
21 September 2006, W.Y. Zhuang, J. Luo, W.Y. Li 6846 (HMAS 183542).
Sequences. ACT (MF669046), ITS (HM054141), LSU (HM042430) and RPB1
(MF669058).
Habitat. On bark and dead twigs.
Distribution. Asia (China).
Notes. Morphologically Rugonectria sinica resembles R. castaneicola (W. Yamam.
& Oyasu) Hirooka & P. Chaverri in having four-spored asci (Zeng et al. 2012). How-
ever, R. castaneicola diers in possessing perithecia that are 250–470 × 350–430 μm
and larger than those of R. sinica that are 216–420 × 194–404 μm. In addition, the
ascospores of R. castaneicola are larger, 18–28 × 7.5–11 μm, while those of R. sinica are
16–26 × 5.5–11 μm. e sequence analyses of the ITS and β-tubulin regions from type
culture conrmed that they are dierent taxa (Zeng et al. 2012).
elonectria guangdongensis Z.Q. Zeng & W.Y. Zhuang, sp. nov.
Fungal Names: FN570488
Figure 3
Holotype. CHINA. Guangdong, Shixing, Chebaling, (24°43'17.38"N,
114°16'39.50"E), alt. 600 m, on branches, 2 November 2015, Z.Q. Zeng, X.C. Wang,
K. Chen, Y.B. Zhang 10627 (HMAS 254522); ex-type culture: HMAS 247233.
Sequences. ACT (MF669045), ITS (MF669051), LSU (MF669053) and RPB1
(MF669057).
Etymology. e specic epithet refers to the type locality of the fungus.
Description. Mycelium not visible around ascomata or on natural substrata. Asco-
mata perithecial, solitary to gregarious, up to 10 in a group, with a well–developed stroma,
supercial, subglobose to globose, bright red with a prominently darkened papilla, turn-
ing dark red in KOH, becoming slightly yellow in LA, 235–382 × 245–412 μm (n =
8). Perithecial surface slightly roughened. Perithecial wall of two layers, 20–50 μm thick,
outer layer 13–37 μm thick, of textura intricata; inner layer 7.5–13 μm thick, of textura
prismatica. Asci not observed. Ascospores ellipsoid, 1-septate, smooth, 10–13 × 3–5 μm
(11.6 ± 1.3 × 4.2 ± 0.7 μm). Colony on PDA 28 mm diameter after 7 d under daylight at
25 °C, surface velvety, with white aerial mycelium, producing purple pigment in medium.
Colony on SNA 35 mm diameter after 7 d under daylight at 25°C, surface with sparse
whitish aerial mycelium. Phialides cylindrical or slightly swollen, 20–58 × 2–4 μm. Macro-
conidia cylindrical, slightly curved with rounded ends, 2–5-septate, 48–70 × 4.8–5.3 μm
(58.9 ± 7.14 × 5.0 ± 0.2 μm). Microconidia and chlamydospores not observed in culture.
Habitat. On branches.
Distribution. Asia (China).
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
110
Figure 3. elonectria guangdongensis a–d ascomata on natural substratum ecolony on PDA f colony on
SNA g median section through perithecium h–m ascospores n , q , r conidiogenous cells and macroco-
nidia o, p , s – u macroconidia. Scale bars: 0.5 mm (a–d); 50 μm (g); 10 μm (h–u).
Notes. Amongst species of elonectria, T. guangdongensis resembles T.phoenicea
in having subglobose to globose perithecia with slightly roughened surface, purple
colony, lack of microconidia and number of septa in macroconidia (Salgado-Salazar
et al. 2015). However, T. phoenicea has much larger perithecia 300–600 × 200–350
e genera Rugonectria and elonectria in China 111
μm, wider ascospores that are 4–5.5 μm wide, and wider phialides 3–6.5 μm wide
(Salgado-Salazar et al. 2015). Moreover, there are 13 bp, 44 bp, 8 bp and 54 bp
divergences in the ACT, ITS, LSU and RPB1 regions, respectively, between the type
of T. guangdongensis (HMAS 254522) and that of T.phoenicea (G.J.S. 85–179).
Phylogenetically T. guangdongensis is closely related to T. beijingensis with strong
statistical support (MLBP/MPBP/BIPP = 100%/97%/100%) (Figure 1). However, T.
beijingensis diers in having larger ascospores that are 13–17 × 4–7 μm, while those of
T. guangdongensis are 10–13 × 3–5 μm and form microconidia in culture in addition
to macroconidia (Zeng and Zhuang 2013). ere are 20 bp, 30 bp, 5 bp and 50 bp
divergences in the ACT, ITS, LSU and RPB1 regions between the ex-type culture of
T. guangdongensis and that of T. beijingensis (HMAS 188498). Both morphology and
molecular data support the establishment of the new species.
elonectria beijingensis Z.Q. Zeng, J. Luo & W.Y. Zhuang, Phytotaxa 85(1): 18, 2013
Specimen examined. CHINA. Beijing, on bark of an unidentied tree, 1 September
2010, L. Cai 7604 (HMAS 188498), ex-type culture: HMAS 188566.
Sequences. ACT (MF669047), ITS (JQ836656), LSU (MF669054) and RPB1
(MF669059).
Habitat. On bark.
Distribution. Asia (China).
Notes. is species was introduced by Zeng and Zhuang (2013) and only known
from the type locality. e phylogenetic analyses indicate that the species is associated
with T. guangdongensis (Figure 1).
elonectria coronalis C. Salgado & Guu, in Salgado-Salazar, Rossman, Samuels,
Capdet & Chaverri, Mycologia 104(6): 1339, 2012
Habitat. On bark of decaying shrubs and trees.
Distribution. Asia (China).
Notes. Salgado-Salazar et al. (2012) described T. coronalis, based on the specimens
occurring on bark of decaying shrubs and trees. e fungus is only known from Taipei
and Yilan of Taiwan Province.
elonectria coronata (Penz. & Sacc.) P. Chaverri & C. Salgado, in Chaverri, Sal-
gado, Hirooka, Rossman & Samuels, Stud. Mycol. 68: 76, 2011
≡ Nectria coronata Penz. & Sacc., Malpighia 11(11–12): 510, 1897.
Specimen examined. CHINA. Hainan, Lingshui, Diaoluoshan, alt. 1050 m, on rotten
twigs of Pinus sp., 15 December 2000, W.Y. Zhuang, X.M. Zhang H90 (HMAS 76855).
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
112
Habitat. On bark of shrubs and trees, sometimes associated with small cankers.
Distribution. Americas (Costa Rica), Asia (Indonesia, Taiwan), possibly pan-
tropical.
Notes. e morphology and molecular data indicated that T. coronata is a species
complex. Salgado-Salazar et al. (2012) divided it into ve taxa on the basis of multi-
gene phylogeny. e Chinese collection matches well the concept of T. coronata sensu
stricto by Salgado-Salazar et al. (2012).
elonectria discophora (Mont.) P. Chaverri & C. Salgado, in Chaverri, Salgado,
Hirooka, Rossman & Samuels, Stud. Mycol. 68: 76, 2011
≡ Sphaeria discophora Mont., Annls Sci. Nat., Bot., sér. 2 3: 353, 1835.
≡ Neonectria discophora (Mont.) Mantiri & Samuels, in Mantiri, Samuels, Rahe &
Honda, Can. J. Bot. 79(3): 339, 2001.
Specimens examined. CHINA. Hainan, Changjiang, Bawangling, alt. 1100 m, 7 De-
cember 2000, on rotten twigs, W.Y. Zhuang, X.M. Zhang, Z.H. Yu H24 (HMAS
83351); Lingshui, Diaoluoshan, alt. 1050 m, 15 December 2000, on rotten twigs,
W.Y. Zhuang, X.M. Zhang H83, H92-1 (HMAS 83353, 83352). Yunnan, Teng-
chong, 16 October 2003, W.P. Wu W7097 (HMAS 183180).
Habitat. On decaying bark of shrubs and trees.
Distribution. Americas (Chile), Asia (China), Europe (Scotland).
Notes. elonectria discophora is the type species of the genus elonectria. Many
specimens identied as this species were determined to be species complex until Sal-
gado-Salazar et al. (2015) separated them into at least 16 taxa, based on phylogenetic
analyses of six nuclear loci and morphological evidences.
elonectria ianthina C. Salgado & Guu, in Salgado-Salazar, Rossman, Samuels,
Hirooka, Sanchez & Chaverri, Fungal Diversity 70(1): 12, 2015
Habitat. On decaying bark of trees and shrubs.
Distribution. Americas (Costa Rica), Asia (China).
Notes. is species is known from Heredia Province of Costa Rica and Taiwan
Province of China on decaying bark of trees and shrubs (Salgado-Salazar et al. 2015).
elonectria japonica C. Salgado & Hirooka, in Salgado-Salazar, Rossman, Samu-
els, Hirooka, Sanchez & Chaverri, Fungal Diversity 70(1): 14, 2015
Specimens examined. CHINA. Hubei, Wufeng, Houhe, alt. 800 m, 13 September
2004, on rotten twigs, W.Y. Zhuang, Y. Nong 5621 (HMAS 98327); Yunnan, Teng-
chong, on rotten twigs, W.P. Wu W7104a (HMAS 183155).
e genera Rugonectria and elonectria in China 113
Sequences. ACT (MK556799), ITS (HM054140) and LSU (HM042434).
Habitat. On decaying bark of Fagus crenata and possibly on bark of other shrubs
and trees.
Distribution. Asia (China, Japan).
Notes. Specimens of this fungus were treated as T. discophora sensu lato until
T. japonica was introduced by Salgado-Salazar et al. (2015). e morphological
characteristics of the Chinese materials t the concept of T. japonica. e Hubei and
Yunnan collections extend its distribution to China.
elonectria lucida (Höhn.) P. Chaverri & C. Salgado, in Chaverri, Salgado, Hi-
rooka, Rossman & Samuels, Stud. Mycol. 68: 76, 2011
≡ Nectria lucida Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 298,
1909.
≡ Neonectria lucida (Höhn.) Samuels & Brayford, in Brayford, Honda, Mantiri &
Samuels, Mycologia 96(3): 590, 2004.
Habitat. On decaying bark of shrubs and trees.
Distribution. Africa (Cameroon), Americas (Costa Rica), Asia (China, Indone-
sia), possibly pantropical.
Notes. is is a relatively common species and recorded as Neonectria lucida by
Guu et al. (2007) from Taiwan Province.
elonectria mamma C. Salgado & P. Chaverri, in Salgado-Salazar, Rossman &
Chaverri, Fungal Diversity 80: 444, 2016
Habitat. On decaying bark of shrubs and trees.
Distribution. Americas (French Guiana), Asia (China).
Notes. e specimens of this species were led under T. lucida (Guu et al. 2007).
After re-examinations of the collections from China and French Guiana, Salgado-Sa-
lazar et al. (2016) stated that they represent a separate species related to T. discophora
sensu stricto.
elonectria phoenicea C. Salgado & P. Chaverri, in Salgado-Salazar, Rossman,
Samuels, Hirooka, Sanchez & Chaverri, Fungal Diversity 70(1): 16, 2015
Specimen examined. CHINA. Hainan, Lingshui, Diaoluoshan, alt. 1050 m, 15 De-
cember 2000, W.Y. Zhuang, X.M. Zhang H86 (HMAS 76856).
Sequences. ACT (MK556800), ITS (JQ836657) and LSU (DQ119572).
Habitat. On decaying Acacia celsa and other plants.
Distribution. Asia (China, Indonesia), Oceania (Australia).
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
114
Notes. Re-examination of HMAS 76856 indicated that T. phoenicea is the correct
name for the specimen which was previously identied as T. discophora. It is distributed
also in Taiwan Province (Salgado-Salazar et al. 2015).
elonectria porphyria C. Salgado & Hirooka, in Salgado-Salazar, Rossman, Sam-
uels, Hirooka, Sanchez & Chaverri, Fungal Diversity 70(1): 19, 2015
Specimen examined. CHINA. Hubei, Wufeng, Houhe, alt. 800 m, on rotten twigs,
12 September 2004, W.Y. Zhuang, Y. Nong 5542 (HMAS 98333).
Sequences. ACT (MK556798), ITS (HM054136) and LSU (HM042433).
Habitat. On decaying bark of Cryptomeria japonica and other woody substrates.
Distribution. Asia (China, Japan).
Notes. e collection was previously treated as T. discophora sensu lato (Zhuang
2013). e sequence analyses (Figure 1) and morphological characteristics of HMAS
98333 indicate that the correct name for the collection is T. porphyria.
elonectria sinensis (J. Luo & W.Y. Zhuang) Z.Q. Zeng & W.Y. Zhuang, Phyto-
taxa 85(1): 18, 2013
≡ Neonectria sinensis J. Luo & W.Y. Zhuang, Mycologia 102(1): 147, 2010.
Specimen examined. CHINA. Hubei, Shennongjia, alt. 1700 m, on bark of a conif-
erous (?) tree, 17 September 2003, X.M. Zhang, Y.Z. Wang Z108 (HMAS 183186),
ex-type culture: HMAS 173255.
Sequences. ACT (MF669048), ITS (FJ560441), LSU (FJ560436) and RPB1
(MF669060).
Habitat. On bark of a coniferous (?) tree.
Distribution. Asia (China).
Notes. e species was originally placed in Neonectria by Luo and Zhuang (2010).
e anatomic structures and DNA data support its placement in elonectria (Zeng
and Zhuang 2013).
elonectria veuillotiana (Sacc. & Roum.) P. Chaverri & C. Salgado, Stud. Mycol.
68: 77, 2011
≡ Nectria veuillotiana Sacc. & Roum., Rev. Mycol. 2: 189, 1880.
≡ Neonectria veuillotiana (Sacc. & Roum.) Mantiri & Samuels, Canda. J. Bot. 79: 339,
2001.
Specimens examined. CHINA. Anhui, Jinzhai, Tiantangzhai, alt. 1000 m, on bark,
24 August 2011, W.Y. Zhuang, H.D. Zheng, Z.Q. Zeng, S.L. Chen 7869 (HMAS
e genera Rugonectria and elonectria in China 115
266577). Hubei, Shennongjia, alt. 1200 m, on rotten twigs associated with other fun-
gi, 15 September 2004, W.Y. Zhuang, Y. Nong 5686 (HMAS 98332); Shennongjia,
alt. 1700 m, on bark associated with other fungi, 15 September 2003, X.M. Zhang, Y.
Z. Wang Z196 (HMAS 183188); Xingshan, Longmenhe, alt. 1800 m, on rotten twigs
associated with other fungi, 18 September 2004, W.Y. Zhuang, Y. Nong 5832 (HMAS
99207). Jilin, Changbaishan, alt. 800 m, on rotten twigs, 27 July 2012, T. Bau, W.Y.
Zhuang, H.D. Zheng, Z.Q. Zeng, Z.X. Zhu, F. Ren 8246 (HMAS 266579); Jiaohe,
Qianjin forest farm, alt. 450 m, on rotten twigs, 23 July 2012, T. Bau, W.Y. Zhuang,
Z.Q. Zeng, H.D. Zheng, Z.X. Zhu, F. Ren 8087b (HMAS 266578). Yunnan, Teng-
chong, on rotten twigs associated with other fungi, 16 September 2003, W.P. Wu
W7095 (HMAS 183568).
Sequences. ITS (HM054151) and LSU (HM042437).
Habitat. On bark of deciduous trees, Eucalyptus sp., Fagus sp., Gleditschiatriacanthos,
Salix sp.
Distribution. Asia (China), Europe (France and Germany), Azores Islands.
Notes. e species was rst placed in Nectria, then in Neonectria (Mantiri et al.
2001) and recently transferred to elonectria by Chaverri et al. (2011). It occurs on
bark of recently killed trees, rarely on wood or leaves and is cosmopolitan in distribu-
tion (Brayford and Samuels 1993; Zhuang 2013).
elonectria yunnanica Z.Q. Zeng & W.Y. Zhuang, Phytotaxa 85(1): 19, 2013
Specimen examined. CHINA. Yunnan, Baoshan, on bark of an unidentied tree, 15
October 2003, W.P. Wu W7122 (HMAS 183564), ex-type culture: HMAS 188567.
Sequences. ACT (MF669049), ITS (FJ560438), LSU (MF669055) and RPB1
(MF669061).
Habitat. On bark.
Distribution. Asia (China).
Notes. elonectria yunnanica is only known from the type locality. It is phylo-
genetically related to T. ostrina (Figure 1). However, T. ostrina has a perithecial wall
25–40 μm while those of T. yunnanica are thicker 49–71 μm and have asci that are
(56–)67–86(−98) × 7–12 μm while those of T. yunnanica are larger, 87–120 × 8.2–9.6
μm. Unlike T. yunnanica, T. ostrina does not forming microconidia in culture (Zeng
and Zhuang 2013; Salgado-Salazar et al. 2015).
Excluded species
elonectria jungneri (Henn.) P. Chaverri & C. Salgado, in Chaverri, Salgado,
Hirooka, Rossman & Samuels, Stud. Mycol. 68: 76, 2011
≡ Nectria jungneri Henn., Bot. Jb. 22: 75, 1895.
≡ Neonectria jungneri (Henn.) Samuels & Brayford, Mycologia 96(3): 580, 2004.
Zhao-Qing Zeng & Wen-Ying Zhuang / MycoKeys 55: 101–120 (2019)
116
≡ Macronectria jungneri (Henn.) C. Salgado & P. Chaverri, in Salgado-Salazar, Ross-
man & Chaverri, Fungal Diversity 80: 448, 2016.
Specimen examined. CHINA. Guangdong, Dinghushan, on rotten twigs associated
with other fungi, 9 October 1998, W.P. Wu W1871-2 (HMAS 183155).
Habitat. On various woody substrates, as well as other plant organic matter.
Distribution. Africa (Cameroon), Americas (Brazil, Costa Rica), Asia (China),
possibly pantropical.
Notes. is fungus was originally described as Nectria jungneri and was transferred
to Neonectria (Brayford et al. 2004) and elonectria (Chaverri et al. 2011). e recent
work by Salgado-Salazar et al. (2016) indicated that it belongs to a separate genus
Macronectria C. Salgado & P. Chaverri.
Discussion
e genus Rugonectria is characterised by the non-papillate, orange to red, conspicu-
ously warted to rugose perithecial surface (Chaverri et al. 2011). e ascomatal anat-
omy, perithecial wall reactions to KOH and LA, features of asci and ascospores and
asexual states indicate the placement of R. microconidia in this genus. e multi-locus
sequence analyses conrm our morphological observations (Figure 1) and it is here
described as a new species.
Historically, the nectriaceous fungi with cylindrocarpon -like asexual states were as-
signed to Neonectria. e accumulated morphological and phylogenetic data suggest
that the genus was heterogeneous (Mantiri et al. 2001). Eorts were made towards
establishment of a monophyletic Neonectria as well as its allies (Booth 1966, Rossman
et al. 1999; Mantiri et al. 2001; Brayford et al. 2004). e previously recognised infra-
generic groups within Neonectria are now recognised as separate genera, i.e. Ilyonectria
for the N. radicicola-group, Neonectria sensu stricto for the N. coccinea-group, Rugon-
ectria for the N. rugulosa-group and elonectria for the N. mammoidea/N. veuilloti-
ana-groups (Chaverri et al. 2011). Since the establishment of elonectria, 45 species
have been placed in the genus (www.indexfungorum.org). Salgado-Salazar et al. (2012,
2015) suggested that the criteria formerly used for generic dierentiation were of insuf-
cient sensitivity to accurately reect the degree of species diversity within the group.
Subsequently, Salgado-Salazar et al. (2016) emended the generic concept of elonectria
by excluding T. jungneri, based on the molecular data and morphological characteristics.
e type species of elonectria, T. discophora, previously considered to be cosmo-
politan, was rst described based on material collected from Chile and was determined
to be heterogeneous (Brayford et al. 2004). Salgado-Salazar et al. (2015) provided a revi-
sionary treatment of the T. discophora species complex and recognised 16 cryptic species
on the basis of the combined analyses of phylogeny and morphology. In this study, the
new species T. guangdongensis is determined to be congeneric with T. discophora, while
both the molecular data and morphological characteristics indicate that T. guangdongenis
e genera Rugonectria and elonectria in China 117
is distinct from other species of elonectria. To date, 11 species of elonectria have
been recorded from China (Teng 1936; Salgado-Salazar et al. 2012, 2015, 2016; Zeng
and Zhuang 2012; Zhuang 2013). China is extremely diverse in its climate, vegetation,
geographic structures and multiple niches. Our understanding of species diversity of the
nectriaceous fungi will be signicantly broadened in the near future.
Acknowledgements
e authors would like to thank Dr. A.Y. Rossman for her valuable comments and cor-
rections and Drs. X.C. Wang, K. Chen and Y.B. Zhang for collecting specimens jointly
for this study. is work was supported by the National Natural Science Foundation of
China (nos. 31750001, 31570018, 31870012) and Frontier Key Program of Chinese
Academy of Sciences (No. QYZDY-SSW-SMC029).
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