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The species of
Coniolariella
Rasoul Zare
1
Bita Asgari
Department of Botany, Iranian Research Institute of
Plant Protection, P.O. Box 1454, Tehran 19395, Iran
Walter Gams
Molenweg 15, 3743CK Baarn, the Netherlands
Abstract
:Morphological and molecular analyses
demonstrate that
Coniolariella gamsii
and
Coniolaria
murandii
are distinct species. The latter species is
validated here as
Coniolariella macrothecia
. A key to
the five species of the genus is provided.
Key words:
epiphytic fungi,
Nodulisporium
,tax-
onomy, Xylariaceae
Coniolariella
D. Garcı´a, Stchigel & Guarro 2006,
recently segregated from
Coniochaeta
(Coniochae-
tales), was introduced for species with affinities to
Xylariales (Garcı´a et al. 2006) and replaced the
previous invalidly published (ICBN Art 36.1) name
Coniolaria
Seigle-Mur., Guiraud, Steiman & Sage
1995, typified by
C. murandii
(Seigle-Murandi et al.
1995). The monotype was
C. gamsii
(Asgari & Zare)
Garcı´a, Stchigel & Guarro, with
Coniolaria murandii
as its purported synonym.
Coniolariella
accommo-
dates species having asci lacking visible apical
structures and polyblastic conidiogenesis. In a further
molecular study Checa et al. (2008) added the new
species
C. hispanica
and the new combination
C.
limoniispora
, under which they recognized
C. gamsii
and
Rosellinia australis
Sacc. & Trotter as varieties. In
the meantime Asgari et al. (2007) introduced another
related species,
Coniochaeta ershadii
, without molecu-
lar analysis. In the two molecular studies only an ex-
type strain (LCP 04.5004) of
Coniolaria murandii
,
identified as
C. gamsii
, was used but no authentic
isolates provided by Asgari et al. (2007). It appeared
doubtful that the synonymy of
Coniolaria murandii
and
Coniolariella gamsii
was correct and re-examina-
tion of type material of both taxa confirmed this
suspicion. Because the study by Checa et al. (2008)
was based only on LSU sequences an attempt to
obtain a living culture of
C. limoniispora
was made to
include it in our ITS tree (FIG. 2), but this was not
successful. Therefore the conclusions are based only
on LSU sequences (FIG. 1).
The sequences were aligned with the pairwise
alignment option in GeneDoc (Nicholas and Nicho-
las 1997). Three species described by Asgari and
Zare (2006) and Asgari et al. (2007) in addition to
others from GenBank (TABLE I) were included in
the analyses of partial LSU (D1–D2, 393–397 bp)
and ITS (444–515 bp) phylogeny with parsimony
and neighbor joining analyses in MEGA4 (Tamura
et al. 2007, see also Asgari and Zare 2010).
(Neighbor joining cladograms based on partial
LSU and ITS sequences are shown in FIGS 1, 2.)
Sequences were submitted to GenBank (TABLE I)
and alignments to TreeBASE (SN4942).
Coniolaria
murandii
is validated in
Coniolariella
as
C. macro-
thecia
. Five species are now recognized in
Coniolar-
iella
as follows.
Coniolariella gamsii (Asgari & Zare) Garcı´a, Stchigel
& Guarro (2006: 1285), holotype of
Coniolariella
.
Basionym:
Coniochaeta gamsii
Asgari & Zare, Nova
Hedwigia 82:228 (2006).
Coniolariella ershadii (Zare, Asgari & W. Gams) Zare,
Asgari & W. Gams, comb. nov.
Mycobank MB516539.
Basionym:
Coniochaeta ershadii
Zare, Asgari & W. Gams,
Nova Hedwigia 84:177 (2007).
Coniolariella hispanica Checa, Arenal & J.D. Rogers
(2008: 797).
Coniolariella limoniispora (Ellis & Everh.) Checa,
Arenal & J.D. Rogers (2008: 797).
Basionym:
Rosellinia limoniispora
Ellis & Everh.,
Proc. Acad. Nat. Sci. Phila. 46:326 (1894).
5
Coniolariella limoniispora
var.
australis
Checa, Arenal &
J.D. Rogers (2008: 797), based on
Rosellinia australis
Sacc. & Trotter 1913 [non Speg. 1909].
Coniolariella macrothecia Zare, Asgari & W. Gams, sp.
nov. FIG.3
MycoBank MB516540.
5
Coniolaria murandii
Seigle-Mur., Guiraud, Steiman &
Sage, Crypt. Bot. 5:347 (1995); nom. inval., Art. 36.1.
Coniolariellae gamsii
similis sed ascomatibus majoribus
(vulgo 800–1200 mm diam) crassitunicatis distin-
guenda. Anamorphe
Nodulisporii
similis, conidiis 7–12
32–2.5 mm.
Colonies fast-growing, floccose, plane, with low and
white aerial mycelium, soon becoming pale gray with
Mycologia myco-102-06-13.3d 22/9/10 01:25:29 1383 Cust #10-035R
Submitted 14 Feb 2010; accepted for publication 10 Apr 2010.
1
Corresponding author. E-mail: simplicillium@yahoo.com
Mycologia,
102(6), 2010, pp. 1383–1388. DOI: 10.3852/10-035
#2010 by The Mycological Society of America, Lawrence, KS 66044-8897
1383
the production of ascomata, reverse pale buff;
ascomata superficial, globose, solitary or in small
groups, dark brown to black, 800–1200(–1500) mm,
with short ostioles, up to 50 mm long and 100 mm wide;
peridium 170–250 mm thick, pseudoparenchymatous,
two-layered, outer part of several layers of dark brown,
thick-walled, anastomosing and interwoven hyphae
(
textura intricata
), inner part of several layers of
hyaline to pale olive-brown, thin-walled, subglobose to
angular cells, 5–15 mmdiam(
textura epidermoidea
or
angularis
); paraphyses numerous, filiform, septate,
simple, slightly undulate, considerably longer than
the asci (150–230 mm), 7–10 mm wide at the base,
tapering to 1.5–2.5 mm at the apex; asci cylindrical,
slightly tapering toward the base, eight-spored, 100–
130 312–16 mm; ascospores exuded as a large,
globose to irregularly shaped mass at the ostiole,
obliquely uniseriate in the ascus, hyaline at first, soon
turning dark pistachio green and finally dark choco-
late brown and black, ellipsoidal fusoid, with a
protrusion at one or both ends, 16–20 39–12 mm;
immature ascospores with a distinct large guttule
when mounted in water; germ slit straight to slightly
sinuous at one end (FIG. 3f, g, k–l). Octahedral
crystals present in the culture (FIG. 3u, v).
Holotype: IRAN 14439 F (ex-type strain LCP
04.5004, IRAN 1637 C, CBS 125772), isolated from
soil at Monument Valley area, Colorado, USA, by F.
Seigle-Murandi, 1995.
Anamorph: nodulisporium-like.
Mycologia myco-102-06-13.3d 22/9/10 01:26:20 1384 Cust #10-035R
FIG. 1. NJ tree based on partial LSU-rDNA (D1–D2) sequences. Bootstrap values .50%(1000 replicates) shown above
branches (those of parsimony analysis are shown in brackets below branches).
Coniocessia nodulisporioides
is outgroup.
Col.
5
Coniolariella
,
Con.
5
Coniochaeta
,
X.
5
Xylaria
.
1384 MYCOLOGIA
Conidiophores produced predominantly on asco-
matal initials, variable in length, (15–)25–85(–120)
32–3 mm, macronematous, mononematous, mostly
simple (rarely branched), smooth, septate, hyaline
or pale olive-brown; conidiogenous cells monoblas-
tic or polyblastic, integrated, terminal or intercalary,
with sympodial elongation, visible as short, less than
1mm long, inconspicuous denticles bearing blasto-
conidia solitarily and leaving a distinctly pigmented
scar; conidia holoblastic, hyaline to pale olive,
smooth, oblong, cylindrical with rounded apex and
attenuated and truncated base, 7–12 32–2.5 mm
(FIG.3m–t).
Remarks.
This species is distinguished from
C.
gamsii
by having larger ascomata with a much thicker
peridium. In the absence of information relating to the
etymology of the original epithet
murandii
a different,
descriptive name for this taxon is introduced based on
the largest ascomata thus far known in the genus.
FIVE SPECIES OF
C
ONIOLARIELLA
ARE TO BE DISTINGUISHED AS FOLLOWS:
1. Ascomata non-ostiolate; paraphyses absent in
mature ascomata; ascospores ellipsoidal to slightly
inaequilateral without protruding ends, 16–17.5 3
9–9.5 mm .....................
C. hispanica
19. Ascomata ostiolate; paraphyses present in mature
ascomata; ascospores with protruding ends ..... 2
2. Ascomata 200–500 mm diam, covered with blunt-
ended setae; peridium 100–120 mm thick; asci 110–
150 310.5–12 mm; ascospores 16–18 39.5–
10.5 mm, with distinctive protruding ends up to
1.7–2.5 mm long ..................
C. ershadii
29. Ascomata larger, without setae; asci slightly shorter
(,130 mm); ascospores on average wider with
slightly protruding ends . . ................. 3
3. Ascomata globose to ovate, often flattened or
depressed above, smooth or granular—roughened;
peridium 80–110 mm thick; ascospores 16–18 3
10.5–12 mm; anamorph sporothrix-like . ......
............................
C. limoniispora
39. Ascomata globose to subglobose, not flattened or
depressed above, smooth; anamorph nodulispor-
ium-like ............................... 4
4. Ascomata 500–800 mm diam; peridium 80–110 mm
thick; asci 110–130 312–14 mm; ascospores 16–19
36–11 mm .......................
C. gamsii
49. Ascomata 800–1200 mm diam; peridium 170–
250 mm thick; asci 100–130 312–16 mm; ascospores
16–20 39–12 mm ...............
C. macrothecia
Mycologia myco-102-06-13.3d 22/9/10 01:26:28 1385 Cust #10-035R
FIG. 2. NJ tree based on ITS (rDNA) sequences. Bootstrap values .50%(1000 replicates) shown above branches (those of
parsimony analysis are shown in brackets below branches).
Coniochaeta velutinosa
is outgroup.
Col.
5
Coniolariella
,
R.
5
Rosellinia
.
ZARE ET AL.: SPECIES OF
C
ONIOLARIELLA
1385
Mycologia myco-102-06-13.3d 22/9/10 01:26:33 1386 Cust #10-035R
TABLE I. Isolates used in phylogenetic analyses
Species Strain Source
Sequences
LSU ITS
Col. ershadii
Co48, IRAN 972C, CBS
119785 (T)
Twig of
Pistacia vera
, Iran GU553331 GU553328
Col. gamsii
Co27, IRAN 842C, CBS
114379 (T)
Leaf of
Hordeum vulgare
, Iran GU553329 GU553325
Col. gamsii
Co28, IRAN 907C, CBS
117677
Leaf of
Hordeum vulgare
, Iran — GU553326
Col. hispanica
Co125, ATCC MYA4453,
CBS 124506, TAH 34624
(T)
Leaf of
Eryngium campestre
, Spain EF489465 GU553323
Col. limoniispora
08fa (R108)
Vitis vinifera
, USA EF489469 —
Col. limoniispora
var.
australis
AH 24323 (R154AH) Twig of
Nicotiana glauca
, Spain EF489464 AY908997
Col. macrothecia
Co127, LCP 04.5004,
IRAN 1637 C, CBS
125772 (T)
Soil, USA AJ875233 GU553324
Con. boothii
CBS 381.74 (T) Soil, India AJ875226 —
Con. cruciata
FMR 7409 Soil, Nigeria AJ875222 —
Con. discoidea
CBS 158.80 (T) Soil, Japan AJ875230 —
Con. hansenii
CBS 885.68 Dung of rabbit, the Netherlands AJ875223 —
Con. ornata
FMR7415 Soil, Russia AJ875228 —
Con. ostrea
CBS 507.70 Twig of
Larrea
sp., USA AJ875227 —
Con. punctulata
FMR 7408 Soil, Argentina AJ875231 —
Con. savoryi
CBS 725.74 (T) Wood of
Juniperus scopulorum
, UK AJ875229 —
Con. tetraspora
FMR 8167 Soil, Spain AJ875225 —
Con. velutinosa
Co29, IRAN 843C, CBS
117678 (T)
Leaf of
Hordeum vulgare
, Iran GU553330 GU553327
Con. verticillata
CBS 816.71 (T) Soil, the Netherlands AJ875232 —
Coniocessia
nodulisporioides
CBS 281.77 (T) Soil, Jordan AJ875224 —
Daldinia fissa
35524C
Betula
sp., Finland — AF176979
Nodulisporium
sp. Et0502
Epichloe¨ typhina
on
Anthoxanthum
odoratum
— EF600033
Nodulisporium
sp. JP807 ? — AF280629
R. arcuata
CBS 347.29 Japan? — AB017660
R. buxi
ATCC 32869 ? — AY909000
R. buxi
R153A ? — DQ272572
R. necatrix
W539 A dead broad-leaf tree, Japan — AB430453
X. badia
BCC 1190 Thailand? AB376711 —
X. escharoidea
BCC 23634, NBRC
104736
Soil, Thailand AB376822 —
X. hypoxylon
GB6391 ? AY327481 —
X. mellissii
BCC 1186 Wood, Thailand AB376710 —
X. psidii
BCC 1127 Thailand? AB376698 —
Underlined sequence numbers are generated in this study, others are from GenBank.
Col.
5
Coniolariella
,
Con.
5
Coniochaeta
,
R.
5
Rosellinia
,
X.
5
Xylaria
. (T), ex-type strain; ATCC, American Type Culture Collection, USA; CBS,
Centraalbureau voor Schimmelcultures, Utrecht, the Netherlands; LCP, Laboratoire de Cryptogamie, Paris, France; IRAN,
Herbarium Ministerii Iranici Agriculturae, Tehran, Iran; BCC, BIOTEC Culture Collection, Thailand Biodiversity Center;
others are not registered abbreviations.
1386 MYCOLOGIA
Mycologia myco-102-06-13.3d 22/9/10 01:26:34 1387 Cust #10-035R
FIG.3.
Coniolariella macrothecia
. Teleomorph: a, b. ascomata and masses of ascospores (top view); c. section of ascoma
showing two-layered peridium; d. peridium outer layer; e. peridium inner layer; f, g. immature asci containing ascospores with
distinct guttule; h. mature ascus; i, j. paraphyses; k. mature ascospores with protruding ends; l. germ slits. Anamorph: m–t.
conidiophores, conidiogenous cells and conidia; u, v. crystals. All mounts were prepared in water. Bars: a 51000 mm; b 5
200 mm; c 550 mm; j 520 mm; d–i, k–v 510 mm; f, r–v 5DIC.
ZARE ET AL.: SPECIES OF
C
ONIOLARIELLA
1387
ACKNOWLEDGMENTS
This study was supported in part by a research grant from
Iranian National Science Foundation (INSF). The authors
are indebted to Dr Joe¨lle Dupont (France) for supplying the
ex-type culture (LCP 04.5004) of
Coniolaria murandii
and
to Dr Gerard Verkley (the Netherlands) for supplying the
ex-type culture (CBS 124506) of
Coniolariella hispanica
.
LITERATURE CITED
Asgari B, Zare R. 2006. Two new
Coniochaeta
species from
Iran. Nova Hedwig 82:227–236.
———, ———. 2010. Two new species of
Preussia
from
Iran. Nova Hedwig 90:533–548.
———, ———, Gams W. 2007.
Coniochaeta ershadii
, a new
species from Iran, and a key to well documented
Coniochaeta
species. Nova Hedwigia 84:175–187.
Checa J, Arenal F, Blanco N, Rogers JD. 2008.
Coniolariella
hispanica
sp. nov. and other additions to
Coniolariella
.
Mycol Res 112:795–801.
Garcı´a D, Stchigel AM, Cano J, Calduch M, Hawksworth DL,
Guarro J. 2006. Molecular phylogeny of
Coniochaetales
.
Mycol Res 110:1271–1289.
Nicholas KB, Nicholas HB Jr. 1997. GeneDoc: a tool for
editing and annotating multiple sequences alignments.
http://www.psc.edu/biomed/genedoc. Distributed by
the authors.
Seigle-Murandi F, Guiraud F, Steiman R, Sage L. 1995.
Coniolaria
: a new genus of Ascomycetes—description of
Coniolaria murandii
sp. nov. from soil of Monument
Valley in Colorado (USA). Crypt Bot 5:346–350.
Tamura K, Dudley J, Nei M, Kumar S. 2007. MEGA4:
molecular evolutionary genetics analysis (MEGA)
software version 4.0. Mol Biol Evol 10.1093/molbev/
msm092.
Mycologia myco-102-06-13.3d 22/9/10 01:27:22 1388 Cust #10-035R
1388 MYCOLOGIA