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Endophragmiella spp. nov. and new records from southern China

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Four new species— Endophragmiella chinensis , E. guangdongensis , E. lushanensis , and E. obovoidea —collected on dead branches of unidentified plants in southern China, are described, illustrated, and compared with closely related species. From the same habitat, three other species, E. cantabrica , E. ellisii , and E. uniseptata are recorded from China for the first time.
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MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2021
January–March 2021—Volume 136, pp. 169–182
https://doi.org/10.5248/136.169
Endophragmiella spp. nov. and
new records from southern China
J M*, L-G M, Z-H X, R-Q C,
L Q, R F. C-R, X-G Z
1College of Agronomy, Jiangxi Agricultural University, Nanchang, Jiangxi 330045, China
2Shandong Key Laboratory of Plant Virology, Institute of Plant Protection,
Shandong Academy of Agricultural Sciences, Jinan, Shandong 250100, China
3Department of Plant Pathology, Shandong Agricultural University,
Taian, Shandong 271018, China
4Instituto de Investigaciones Fundamentales en Agricultura Tropical ‘Alejandro de Humboldt’
(INIFAT), Académico Titular de la “Academia de Ciencias de Cuba”,
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
*C : majian821210@163.com; jxaumj@126.com
A—Four new species—Endophragmiella chin ensis, E. guangdongensis, E. lushanensis,
and E. obovoidea—collected on dead branches of unidentied plants in southern China, are
described, illustrated, and compared with closely related species. From the same habitat,
three other species, E. cantabrica, E. ellisii, and E. uniseptata are recorded from China for
the rst time.
K —asexual Ascomycota, hyphomycetes, saprobes, taxonomy
Introduction
Endophragmiella B. Sutton was established (Sutton 1973) for two species,
E. pallescens B. Sutton (generic type) and E. canadensis (Ellis & Everh.)
B. Sutton [= E. subolivacea (Ellis & Everh.) S. Hughes; see Hughes 1979].
Endophragmiella is mainly characterized by solitary, acrogenous, aseptate
or euseptate conidia seceding rhexolytically from monoblastic, integrated,
terminal, determinate or percurrently elongating conidiogenous cells
on simple or branched conidiophores (Sutton 1973, Ellis 1976, Hughes
170 ... Ma & al.
1979, Holubová-Jechová 1986, Wang 1990, Wu & Zhuang 2005, Seifert &
al. 2011). Hughes (1979) claried the conidiogenesis of Endophragmiella,
amended the generic concepts, and accepted 33 species. Subsequently, 62
species and one variety have been published in Endophragmiella (Index
Fungorum 2020). Endophragmiella cambrensis M.B. Ellis (Ellis 1976)
was excluded from that genus by Hughes (1979) but accepted by Wu
& Zhuang (2005). Endophragmiella dimorphospora Awao & Udagawa
ex P.M. Kirk was treated as a synonym of the prior name E. biconstituta
(Rambelli) Matsush. by Matsushima (1989). Endophragmiella fasciata
(R.F. Castañeda) R.F. Castañeda (Castañeda-Ruíz 1988) and E. quadrilocularis
Matsush. (Matsushima 1993) were transferred to Repetophragma Subram.
by Castañeda-Ruíz & al. (2006, 2011). Endophragmiella fuliginosa
(B. Sutton) S. Hughes (Hughes 1979) was treated as Rhexoacrodictys
fuliginosa (B. Sutton) W.A. Baker & Morgan-Jones by Baker & al. (2002).
Endophragmiella rigidiuscula R.F. Castañeda (Castañeda-Ruiz 1988)
was proposed as the type species of Distophragmia R.F. Castañeda & al.
(Castañeda-Ruíz & al. 2015). Nine invalid Endophragmiella species names
have been validated in Senwanna (2019). Endophragmiella currently
contains 91 species and one variety, of which 46 species have been recorded
from China (Matsushima 1980, Tzean & Chen 1989, Lu & al. 2000, Tsui & al.
2001, Zhuang 2001, Wu & Zhuang 2005, Chen & al. 2008, Zhang & al. 2010,
Ma & al. 2011, 2012a,b, 2015, Ren & al. 2011, Wu & Zhang 2012, Xia & al.
2016, Li & al. 2017, Jiang & al. 2018).
During an ongoing survey of anamorphic fungi on plant debris in
southern China, seven species clearly related to Endophragmiella were
collected from dead branches. Four are described here as new, and the other
three represent new records for the Chinese mycota.
Taxonomy
Endophragmiella chinensis L. Qiu, Jian Ma, R.F. Castañeda & X.G. Zhang,
sp. nov. F. 
IF 
Diers from Endophragmiella bohaniensis by its oblong or cylindrical, smaller conidia
on branched, shorter, and narrower conidiophores; and from E. constricta, E. fallacia,
and E. oblonga by its smaller, concolorous conidia.
T: China, Jiangxi Province, Lushan Mountain, on dead branches of an unidentied
broadleaf tree, 18 October 2016, J. Ma (holotype, HJAUP M0477).
E: refers to China where the fungus was collected.
Four new Endophragmiella species (China) ... 171
F. 1. Endophragmiella chinensis (holotype, HJAUP M0477).
A. Conidiophores, conidiogenous cells, and conidia; B. Conidiophore; C, D. Conidia.
C on the natural substrate euse, brown to dark brown. Mycelium
partly immersed, partly supercial, composed of branched, septate,
pale brown, smooth-walled hyphae. C macronematous,
mononematous, single or aggregated at the base, simple or branched,
erect, straight or exuous, septate, smooth, medium brown to brown, paler
towards the apex, 83–115 × 2.5–4 µm. C  monoblastic,
integrated, terminal, cylindrical, smooth, pale brown, attenuated into
172 ... Ma & al.
narrow and truncate at the apex, with 0–2 percurrent elongations. Conidial
secession rhexolytic. C solitary, acrogenous, smooth, thick-walled,
(1–)2-euseptate, the septum usually becoming darker and forming a band;
when mature oblong or cylindrical, medium brown, (11–)14–21 µm long,
(4.5–)6–7.5 µm diam. in the broadest part; apex rounded; base truncate,
1–2 µm diam., with a distinct 1–1.5 µm long basal frill derived from the
distal end of the conidiogenous cell.
C—Among the Endophragmiella taxa with predominantly
2-euseptate conidia, E. chinensis is most similar to E. bohaniensis N.D.
Sharma (Sharma 1985), E. constricta M.T. Dunn (Dunn 1982), E. fallacia
P.M. Kirk (Kirk 1981a), and E. oblonga Matsush. ex S. Hughes (Hughes 1979)
in conidial shape. However, E. bohaniensis diers by its broadly ellipsoidal
to oblong, larger conidia (18–33 × 8.5–12.5 µm) on unbranched longer and
wider conidiophores (75–300 × 5–6 µm; Sharma 1985); E. constricta diers
by its brown larger conidia (15–23 × 6.9–10.4 µm) with paler lower cell
and constricted middle cell (Dunn 1982); E. oblonga and E. fallacia dier
mainly in their unbranched conidiophores, and larger versicolored conidia
(E. oblonga 19–28 × 9.5–11.5 µm, Hughes 1979; E. fallacia 22–26 × 10–12
µm, Kirk 1981a).
Endophragmiella guangdongensis L. Qiu, Jian Ma, R.F. Castañeda &
X.G. Zhang, sp. nov. F. 
IF 
Diers from Endophragmiella cesatii and E. latispora by its smaller predominantly
2-euseptate conidia; from E. lauri by its versicolored wider conidia; and from E. curvata
by its wider conidia with the longest central or suprabasal cell.
T: China, Guangdong Province: Liuxihe National Forest Park, on dead branches of
an unidentied broadleaf tree, 17 October 2014, J. Ma (holotype, HJAUP M0118).
E: refers to the province in which the fungus was found.
C on the natural substrate euse, brown to dark brown. Mycelium
partly immersed, partly supercial, composed of branched, septate, pale brown,
smooth-walled hyphae. C macronematous, mononematous,
single or aggregated at the base, unbranched, erect, straight or exuous, septate,
smooth, brown to dark brown, up to 530 × 4.5–5.5 µm. C
 monoblastic, integrated, terminal, cylindrical, smooth, brown, with up
to 25 or more percurrent elongations. Conidial secession rhexolytic. C
solitary, acrogenous, fusiform or ellipsoidal, smooth, 2(–3)-euseptate, brown
with the paler end cells, 20.5–29.5 µm long, 8.5–10.5 µm diam. in the broadest
Four new Endophragmiella species (China) ... 173
F. . Endophragmiella guangdongensis (holotype, HJAUP M0118).
A. Conidiophore; B, C. Conidiophores, conidiogenous cells, and conidia; D. Conidia.
part; apex rounded or acute; base truncate, 2.5–4 µm diam., with a distinct
0.5–1 µm long basal frill derived from the distal end of the conidiogenous cell.
C—In terms of conidial morphology, E. guangdongensis is most
similar to E. latispora W.P. Wu (Wu & Zhuang 2005), E. lauri P.M. Kirk &
C.M. Kirk (Kirk 1982), and E. curvata (Corda) S. Hughes and E. cesatii
(Mont.) S. Hughes (Hughes 1979), but E. cesatii and E. latispora dier by
their predominantly 3-euseptate, larger conidia (E. cesatii 32–40(–45) ×
11–12.5 µm, Hughes 1979; E. latispora 23–35 × 12–13 µm, Wu & Zhuang 2005);
E. curvata diers by its narrower (7.2–8.3 µm) conidia with the lowest cell
longer than other cells; and E. lauri diers by its concolorous, narrower
(7–8 µm) conidia (Kirk 1982).
174 ... Ma & al.
F. 3. Endophragmiella lushanensis (holotype, HJAUP M0436).
A–C. Conidiophores, conidiogenous cells, and conidia; D. Conidia.
Endophragmiella lushanensis L. Qiu, Jian Ma, R.F. Castañeda &
X.G. Zhang, sp. nov. F. 
IF 
Diers from Endophragmiella fagicola, E. latispora, E. socia, and E. valdiviana by its
conspicuously smaller predominantly 4-euseptate conidia.
T: China, Jiangxi Province, Lushan Mountain, on dead branches of an unidentied
broadleaf tree, 18 October 2016, J. Ma (holotype, HJAUP M0436).
E: refers to the locality in which the fungus was found.
C on the natural substrate euse, brown to dark brown. Mycelium
partly immersed, partly supercial, composed of branched, septate, pale brown,
smooth-walled hyphae. C macronematous, mononematous,
single or aggregated at the base, unbranched, erect, straight or exuous,
septate, smooth, medium brown to dark brown at the lower part, pale brown
towards the apex, 54–160 × 4.5–6 µm. C  monoblastic,
integrated, terminal, cylindrical, smooth, pale brown, attenuated into narrow
and truncate apex of 2.5–3 µm wide, determinate, with up to 3 percurrent
elongations. Conidial secession rhexolytic. C solitary, acrogenous,
ellipsoidal, fusiform or obclavate, smooth, (3–)4(–5)-euseptate, the septum
Four new Endophragmiella species (China) ... 175
usually becoming darker and forming a band, medium brown to brown, apical
cell pale brown to subhyaline, 20–29 µm long, 7–9 µm diam. in the broadest
part; apex rounded; base truncate, 2.5–3 µm diam., with a distinct 0.5–1 µm
long basal frill derived from the distal end of the conidiogenous cell.
C—Among the known species of Endophragmiella, E. lushanensis is
most similar in conidial shape to E. fagicola P.M. Kirk (Kirk 1981b), E. latispora
(Wu & Zhuang 2005), E. socia (M.B. Ellis) S. Hughes, and E. valdiviana (Speg.)
S. Hughes (Hughes 1979), but E. fagicola diers by its obvious larger (70–90 ×
11–17 µm) usually 5-euseptate conidia (Kirk 1981b); E. latispora diers by its
wider (12–13 µm) predominantly 3-euseptate conidia (Wu & Zhuang 2005);
E. socia and E. valdiviana dier by their larger predominantly 7-euseptate
conidia (E. socia 36–50 × 10.8–14.4 µm; E. valdiviana 40–50 × 11–12.5 µm)
(Hughes 1979).
Endophragmiella obovoidea L. Qiu, Jian Ma, R.F. Castañeda & X.G. Zhang,
sp. nov. F. 
IF 
Diers from Endophragmiella occidentalis, E. aseptata, and E. fatrensis by its obovoid,
smooth, pale brown, smaller conidia.
T: China, Jiangxi Province: Lushan Mountain, on decaying twigs of an unidentied
broadleaf tree, 18 October 2016, J. Ma (holotype, HJAUP M0475).
E: refers to the obovoid conidial shape.
C on the natural substratum euse, brown to dark brown. Mycelium
partly immersed, partly supercial, composed of branched, septate, pale brown,
smooth-walled hyphae. C macronematous, mononematous,
single or aggregated at the base, simple, erect, straight or exuous, septate,
smooth, pale brown to brown, paler towards the apex, ≤230 × 3–4 µm.
C  monoblastic, integrated, terminal, cylindrical, smooth,
pale brown, attenuated into narrow and truncate at the apex, with up to 26 (or
more) percurrent elongations. Conidial secession rhexolytic. C solitary,
acrogenous, smooth, thick-walled, obovoid, sometimes ellipsoidal, aseptate,
pale brown, 8–9.5 × 5–6 µm, with a distinct 0.5–1.5 µm long basal frill derived
from the distal end of the conidiogenous cell.
C—Among the known species of Endophragmiella with aseptate
conidia, E. obovoidea somewhat resembles E. occidentalis R.F. Castañeda & al.
(Castañeda-Ruíz & al. 1995), E. aseptata Hol.-Jech., and E. fatrensis Hol.-Jech.
(Holubová-Jechová 1986), but E. occidentalis diers by its obovate, smooth,
rarely verrucose, dark brown, larger conidia (10–12 × 9–11 µm; Castañeda-
176 ... Ma & al.
F. . Endophragmiella obovoidea (holotype, HJAUP M0475).
A–C. Conidiophores, conidiogenous cells, and conidia; D. Conidia.
Ruíz & al. 1995); E. aseptata diers by its ellipsoidal or ovoid, brown, larger
conidia (7–11 × 6–7.5 µm; Holubová-Jechová 1986); and E. fatrensis diers by
its ellipsoidal to ovoidal, mid to dark smoke-brown, larger conidia (11–15.5 ×
7–8.5 µm; Holubová-Jechová 1986].
Endophragmiella cantabrica J. Mena, Hern.-Restr., Gené & Guarro,
Mycotaxon 123: 225, 2013. F. 
C macronematous, mononematous, single, unbranched,
erect, straight or exuous, 7–13-septate, smooth, brown to dark brown, paler
Four new Endophragmiella species (China) ... 177
F. 5. Endophragmiella cantabrica (HJAUP M001).
A–C. Conidiophores, conidiogenous cells, and conidia; D. Conidia.
towards the apex, ≤220 × 3.5–4.5 µm. C  monoblastic,
integrated, terminal, cylindrical, smooth, brown, with ≤20 percurrent
elongations. Conidial secession rhexolytic. C solitary, acrogenous,
oblong, ellipsoidal or sometimes obovoid, smooth, thick-walled, medially
1-euseptate, pale brown, 13–15.5 µm long, 5.5–7 µm diam. in the broadest part;
apex rounded; base truncate, 2–2.5 µm diam., with a distinct 0.5–1 µm long
basal frill derived from the distal end of the conidiogenous cell.
S : CHINA, J P, Jinggangshan Mountain, on dead
branches of an unidentied broadleaf tree, 16 October 2012, J. Ma (HJAUP M001).
178 ... Ma & al.
F. 6. Endophragmiella ellisii (HJAUP M0316).
A–C. Conidiophores, conidiogenous cells, and conidia; D. Conidia.
C—Hernández-Restrepo & al. (2013) recorded this fungus on dead
wood from Spain, and compared it with similar species, including E. arranensis
P.M. Kirk, E. bogoriensis Rifai, E. ovoidea P.M. Kirk, and E. uniseptata var. pusilla
Hol.-Jech. is is the rst report of E. cantabrica in China. Our collection ts
well with the original description of E. cantabrica except for our slightly larger
conidia (cf. 10–14 × 5–6 µm; Hernández-Restrepo & al. 2013).
Endophragmiella ellisii P.M. Kirk, Index Fungorum 421: 1, 2019. F. 
Endophragmia biseptata M.B. Ellis, Mycol. Pap.72: 31. 1959, nom. inval.
Endophragmiella ellisii” S. Hughes, N.Z. J. Bot. 17(2): 150, 1979, nom. inval.
C macronematous, mononematous, single or aggregated at
the base, unbranched, erect, straight or exuous, 4–9-septate, smooth, brown
to dark brown, paler towards the apex, 70–140 × 5–6.5 µm. C
 monoblastic, integrated, terminal, cylindrical, smooth, pale brown, with
Four new Endophragmiella species (China) ... 179
≤14 percurrent elongations. Conidial secession rhexolytic. C solitary,
acrogenous, obovoid to pyriform, smooth, thick-walled, 2-euseptate, the
septum usually becoming darker and forming a band, distal cell brown to dark
brown, central cell pale brown to brown and basal cell paler, 17.5–23.5 µm long,
8–11.5 µm diam. in the broadest part; apex rounded; base truncate, 2.5–3.5 µm
diam., with a distinct 0.5–2 µm long basal frill derived from the distal end of
the conidiogenous cell.
S : CHINA, J P, Jiulianshan National Nature
Reserve, on dead branches of an unidentied broadleaf tree, 3 November 2014, J. Ma
(HJAUP M0316); H P, Jianfengling National Nature Reserve, on dead
branches of an unidentied broadleaf tree, 15 April 2015, J. Ma (HJAUP M0402).
C –e previous invalid publication of “E. ellisii” was validated in
Senwanna (2019). is species has not previously been recorded from China.
Our specimen overlaps well with the original descriptions of Ellis (1959).
Endophragmiella ellisii is most similar to E. bisbyi B. Sutton ex P.M. Kirk
(Senwanna 2019) and E. hughesii D. Hawksw. (Hawksworth 1979) in conidial
shape, but E. bisbyi diers by its predominantly 3-euseptate, smaller conidia
(12.5–16 × 6.3–7.6 µm; Hughes 1979), and E. hughesii diers by its larger
conidia (25–30 × 11–13 µm) with a central pore at each septum (Hawksworth
1979).
Endophragmiella uniseptata M.B. Ellis ex P.M. Kirk,
Index Fungorum 421: 1, 2019. F.
Endophragmia uniseptata M.B. Ellis, Mycol. Pap.72: 28. 1959, nom. inval.
Endophragmiella uniseptata” M.B. Ellis ex S. Hughes, N.Z. J. Bot. 17(2): 156, 1979, nom.
inval.
C macronematous, mononematous, single or aggregated at the
base, unbranched, erect, straight or exuous, 2–7-septate, smooth, brown to
dark brown, 55–139 × 4–6 µm. C  monoblastic, integrated,
terminal, cylindrical, smooth, pale brown to brown, determinate or with
1–3 percurrent elongations. Conidial secession rhexolytic. C solitary,
acrogenous, obovoid, smooth, 1-euseptate, the septum usually becoming
darker and forming a band, brown to dark brown with the lower cell sometimes
slightly paler, 15.5–21 µm long, 10–12.5 µm diam. in the broadest part; apex
rounded; base truncate, 2.5–3.5 µm diam., with distinct 0.5–1 µm long basal
frill derived from the distal end of the conidiogenous cell.
S : CHINA, G P, Nanling National Nature
Reserve, on dead branches of an unidentied broadleaf tree, 12 July 2014, J. Ma (HJAUP
M0137).
180 ... Ma & al.
F. 7. Endophragmiella uniseptata (HJAUP M0137).
A–C. Conidiophores, conidiogenous cells, and conidia; D, E. Conidiophores; F. Conidia.
C—Ellis (1959) reported this species on rotten wood of Castanea
saliva and Fagus sylvatica from Great Britain. e previous invalid publication
of “E. uniseptata” was validated in Senwanna (2019). Endophragmiella
uniseptata is similar to E. pinicola M.B. Ellis ex P.M. Kirk (Senwanna 2019)
in conidial shape, but E. pinicola has concolorous, smaller conidia (10–14 ×
5–9 µm; Ellis 1976). Our collection corresponds well with the description of
Ellis (1959; as “Endophragmia uniseptata”) except for much fewer percurrent
elongations of conidiogenous cells in our specimen (cf. up to 15; Ellis 1959).
is is the rst report of this species in China.
Acknowledgments
e authors express gratitude to Dr. De-Wei Li (e Connecticut Agricultural
Experiment Station, Valley Laboratory, USA) and Dr. Gabriela Heredia (Instituto
de Ecología, AC, Mexico) for serving as pre-submission reviewers and to Dr. Shaun
Pennycook for nomenclatural review and Dr. Lorelei L. Norvell for editorial review.
is project was supported by the National Natural Science Foundation of China
(Nos. 31970018, 31760513, 31360011).
Four new Endophragmiella species (China) ... 181
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... Comments − Endophragmiella Sutton, is characterised by numerous, successive extensions of the conidiogenous cell and rhexolytic conidial secession, and thin-walled, septate or unicellular conidia (Sutton 1973b (Hughes 1979;Holubová-Jechová 1986;Castañeda-Ruiz et al. 1995;Li et al. 2018;Ma et al. 2021), produce aseptate conidia, and are thus similar to E. pseudostromatica. However, these species vary in conidial colour, shape, and size, and also, E. albiziae (M.B. ...
... They are essential components in forest ecosystems and play important roles in decomposition and nutrient recycling, but their geographical distribution and alpha taxonomy are poorly studied. Our group has investigated saprobic hyphomycetes for about 17 years during which many new genera and species have been named (e.g., Zhang & Shi 2005, Zhang & al. 2009Xia & al. 2015;Ma & al. 2016Ma & al. , 2021Zhang 2018;Xu & al. 2019Xu & al. , 2020. Our continuing survey of saprobic hyphomycetes from dead branches in Guangdong Province, China, revealed a previously undescribed species, proposed here as Zasmidium sinense. ...
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