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Pseudocosmospora, a new genus to accommodate Cosmospora vilior and related species

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Cosmospora sensu Rossman accommodated nectroid fungi with small, reddish, smooth, thin-walled perithecia but recently was found to be polyphyletic and has been segregated into multiple genera. Not all Cosmospora-like fungi have been treated systematically. Some of these species include C. vilior and many specimens often labeled "Cosmospora sp." The objectives of this research were to establish the identity of C. vilior through epitypication using a recent collection that agrees with the type specimen in morphology, host and geography and to determine its phylogenetic position within Cosmospora sensu lato and the Nectriaceae. A multilocus phylogeny was constructed based on six loci (ITS, LSU, MCM7, rpb1, tef1, tub) to estimate a phylogeny. Results from the phylogenetic analyses indicated that C. vilior forms a monophyletic group with other cosmospora-like fungi that have an acremonium-like anamorph and that parasitize Eutypa and Eutypella (Ascomycota, Sordariomycetes, Xylariales, Diatrypaceae). The group is phylogenetically distinct from other previously segregated genera. A new genus, Pseudocosmospora, is described to accommodate the type species, P. eutypellae, and nine additional species in this clade.
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Pseudocosmospora
, a new genus to accommodate
Cosmospora vilior
and related species
Cesar S. Herrera
1
University of Maryland, Department of Plant Science
and Landscape Architecture, 2112 Plant Sciences
Building, College Park, Maryland 20742
Amy Y. Rossman
Gary J. Samuels
United States Department of Agriculture, Agriculture
Research Service, Systematic Mycology and Microbiology
Laboratory, B-010A, 10300 Beltsville, Maryland
20705
Priscila Chaverri
University of Maryland, Department of Plant Science
and Landscape Architecture, 2112 Plant Sciences
Building, College Park, Maryland 20742
Abstract
:
Cosmospora
sensu Rossman accommodated
nectroid fungi with small, reddish, smooth, thin-
walled perithecia but recently was found to be
polyphyletic and has been segregated into multiple
genera. Not all cosmospora-like fungi have been
treated systematically. Some of these species include
C. vilior
and many specimens often labeled ‘‘
Cosmos-
pora
sp.’’ The objectives of this research were to
establish the identity of
C. vilior
through epitypica-
tion using a recent collection that agrees with the type
specimen in morphology, host and geography and to
determine its phylogenetic position within
Cosmos-
pora
sensu lato and the Nectriaceae. A multilocus
phylogeny was constructed based on six loci (ITS,
LSU, MCM7,
rpb1
,
tef1
,
tub
) to estimate a phylogeny.
Results from the phylogenetic analyses indicated that
C. vilior
forms a monophyletic group with other
cosmospora-like fungi that have an acremonium-like
anamorph and that parasitize
Eutypa
and
Eutypella
(Ascomycota, Sordariomycetes, Xylariales, Diatrypa-
ceae). The group is phylogenetically distinct from
other previously segregated genera. A new genus,
Pseudocosmospora
, is described to accommodate the
type species,
P. eutypellae
, and nine additional species
in this clade.
Key words:
fungal systematics, GCPSR, Mycopar-
asite,
Nectria
, one-to-one genus concept
I
NTRODUCTION
Cosmospora
sensu Rossman (Nectriaceae, Hypo-
creales, Ascomycota; Gra¨fenhan et al. 2011) was
erected to accommodate nectroid fungi with small,
reddish, KOH+, smooth, thin-walled, laterally collaps-
ing when dry, non- or weakly stromatic perithecia
(Samuels et al. 1991, Rossman et al. 1999). These
fungi have been reported worldwide, but they are
assumed to have greater diversity in warm temperate
and tropical regions. In addition, they tend to have a
higher diversity in recently disturbed stands (1–2 y
old) compared to early successional stands (25–27 y
old) and old-growth stands in tropical forests (Cha-
verri and ´lchez 2006). In that study, frequently
collected species in recently disturbed stands, where
newly killed woody substrates and herbaceous debris
are prevalent, were members of
Chaetopsinectria
Lou
& Zhuang and
Volutellonectria
Lou & Zhang, two
genera segregated from
Cosmospora
sensu Rossman
(Luo and Zhuang 2010, 2012). Many species of
Cosmospora
sensu Rossman are parasites of their
fungal hosts (see Tsuneda 1982). Among genera
segregated from
Cosmospora
sensu Rossman, some
members of
Cosmospora
sensu stricto grow on
basidiomycetes or xylariaceous hosts; species of
Dialonectria
(Sacc.) Cooke occur on
Diatrype
Fr.
(Diatrypaceae); and
Microcera
Desm. parasitize scale
insects (Gra¨fenhan et al. 2011).
The generic name
Cosmospora
has been a source of
much taxonomic confusion. Rabenhorst (1862) de-
scribed this genus that was reduced to a subgenus of
Nectria
(Fr.) Fr. by Saccardo (1883). Much later, it was
synonymized with
Dialonectria
(Moravec 1954), which
had been elevated from a subgenus of
Nectria
to
generic rank by Cooke (1884). Rossman et al. (1999)
resurrected the generic name
Cosmospora
based on
priority. The group also has been referred to as
Nectria
subgenus
Dialonectria
or the ‘‘
Nectria epi-
sphaeria
-group’’ (Booth 1959, Samuels et al. 1991,
Rossman et al. 1999). Early on, the group was
presumed to be polyphyletic given its range of
anamorphs and ecological niches (Samuels et al.
1991), and at one time as many as 70 species were
classified under
Cosmospora
(www.indexfungorum.
org). The polyphyly of
Cosmospora
was confirmed by
Zhuang and Zhuang (2006), Luo and Zhuang (2008),
Samuels et al. (2009) and Gra¨fenhan et al. (2011).
Following the genus-for-genus concept, that is the
delimitation of a genus based on the correlation of
Submitted 12 Dec 2012; accepted for publication 4 Apr 2013.
1
Corresponding author. E-mail: csherrer@umd.edu
Mycologia,
105(5), 2013, pp. 1287–1305. DOI: 10.3852/12-395
#
2013 by The Mycological Society of America, Lawrence, KS 66044-8897
1287
the teleomorph to its corresponding anamorph
(Rossman 1993),
Cosmospora
was segregated into
new or revived genera that correlate roughly with
these anamorphs:
Chaetopsinectria
,
Cyanonectria
Sa-
muels & P. Chaverri,
Nectricladiella
Crous & Schoch,
Fusicolla
Bonord.,
Macroconia
(Wollenw.) Gra¨fenhan
et al.,
Microcera
,
Stylonectria
Ho¨hn. and
Volutellonec-
tria
(see Schoch et al. 2000; Samuels et al. 2009; Luo
and Zhuang 2010, 2012; Gra¨fenhan et al. 2011). With
the change to one scientific name for each species as
directed in the International Code of Nomenclature
for algae, fungi and plants (ICN) (McNeill et al.
2012),
Chaetopsinectria
and
Volutellonectria
are con-
sidered synonyms of the older genera
Chaetopsina
Rambelli and
Volutella
Fr.
Cosmospora vilior
was described as
Nectria vilior
by
Starba¨ck (1899) with the diagnosis ‘‘Peritheciis
discretis, superficialibus, ovoideis, coccineis…Hab.
in fungillo valsaceo.’’ Traditionally the name has
been applied to collections of cosmospora-like fungi
having short, coarsely warted ascospores occurring on
black stromata, particularly those of the Xylariales
(Weese 1916, Samuels et al. 1990, Samuels et al.
1991).
Nectria vilior
has been reported to have a wide
tropical and temperate distribution (Samuels et al.
1990). Re-examination of the type specimen of
C.
vilior
revealed that its associated host is a species of
Eutypella
(Nitschke) Sacc. (Diatrypaceae). Our recent
molecular analyses suggest that true
C. vilior
is
unrelated to species of
Cosmospora
that occur on
xylariaceous fungi, hereafter referred to as the
C.
viliuscula
species complex. Species of the
C. vilior
complex occur only on species of
Eutypella
(Diatry-
paceae) while
C. viliuscula
and related species are
restricted to xylariaceous fungi.
The current paper deals with the phylogenetic and
taxonomic reassessment of the
Cosmospora vilior
and
similar taxa. The objectives of this research are: (i) to
establish the identity of
C. vilior
and stabilize the
name using epitypification, (ii) to elucidate the
phylogenetic placement of
C. vilior
and related
species within
Cosmospora
sensu Rossman and in the
Nectriaceae, (iii) to describe a new genus,
Pseudocos-
mospora
, to accommodate
C. vilior
and related species
and (iv) to describe new species within
Pseudocosmos-
pora
including the type
P. eutypellae
.
M
ATERIALS AND METHODS
Teleomorph and anamorph morphological characterization.—
Herbarium specimens were borrowed from the U.S.
National Fungus Collections (BPI), the William and Lynda
Steere Herbarium, New York Botanical Garden (NY) and
the Linnean Herbarium, Swedish Museum of Natural
History (S). Fresh specimens were collected on trips to
Argentina, Brazil, Costa Rica, France and USA. For the
characterization of the teleomorph, these observations were
made for perithecia: shape, size (length and width), color,
ornamentation, and habit (e.g. perithecia being solitary or
gregarious, immersed in substrata or superficial, stromatic
or non-stromatic and collapsing laterally or not when dry).
Reaction to 3% w/v potassium hydroxide (KOH) and 100%
lactic acid was observed for the perithecial wall. Sections of
perithecia (ca. 11
mm thick) were made with a freezing
microtome. Measurements of continuous characters (e.g.
length and width) were made with Scion Image software
beta 4.0.2 (Scion Corp., Frederick, Maryland) and summa-
rized by descriptive statistics (e.g. minimum, maximum,
mean and standard deviation).
Cultures were obtained from the culture collection at
USDA, ARS, Systematic Mycology and Microbiology Labo-
ratory (SMML). Additional cultures were obtained by
isolating single ascospores from freshly collected samples
with the aid of a micromanipulator and grown in cornmeal
dextrose agar (CMD; Difco
TM
cornmeal agar + 2% w/v
dextrose + antibiotics). Morphological observations of the
colony were made by growing three pseudoreplicates of
each isolate on CMD and Difco
TM
potato dextrose agar
(PDA) in an incubator that alternated 12 h/12 h between
fluorescent light and darkness at 25 C. Cultural morphology
is described based on strains grown on PDA; cultures on
CMD exhibit little variability. Colony color is described with
the terms in Rayner (1970). Culture growth was measured
weekly for 2 wk. The anamorph was observed by cutting an
agar block of a culture grown in synthetic nutrient-poor
agar (SNA; Nirenberg 1976) under the same conditions
mentioned above, covering it with a cover slip and
examining it by light microscopy (Olympus BX50; Olympus,
Tokyo, Japan). Measurements of continuous characters
were made and analyzed as described above.
DNA extraction, PCR and sequencing.—
The DNA extraction
protocol is described in detail by Hirooka et al. (2010).
Briefly, the isolates were grown in Difco
TM
potato dextrose
broth (PDB) and the mycelial mat was harvested after a
week. DNA was extracted with PowerPlantH DNA Isolation
Kit (MO BIO Laboratories Inc., Solana Beach, California).
Six partial loci were amplified. These loci are internal
transcribed spacer (ITS; primers ITS5, ITS4; White et al.
1990), large subunit nuclear ribosomal DNA (LSU; primers:
LROR and LR5; Vilgalys and Hester 1990), MCM7 (a DNA
replication licensing factor; primers: Mcm7-709 for &
Mcm7-1348 rev; Schmitt el al. 2009), RNA polymerase II
subunit one (
rpb1
; primers: Crpb1a & rpb1c; Castlebury et
al. 2004), translation elongation factor 1-a (
tef1
; primers:
Tef1-728, Tef1-986; Carbone and Kohn 1999) and b-tubulin
(
tub
; O’Donnell and Cigelnik 1997). The PCR reaction
mixture (25
mL total volume) consisted of 12.5 mL Go-
TaqHGreen Master Mix 23 (Promega Corp., Madison,
Wisconsin), 1.25
mL for the forward and reverse primers
each (10 mM), 1.0
mL dimethyl sulfoxide (DMSO; Sigma-
Aldrich, St Louis, Missouri), up to 5.0
mL genomic DNA
template, and RNAse-free water to complete the total
volume. PCR reactions were carried out in an Eppendorf
Mastercycler thermo-cycler (Eppendorf, Westbury, New
1288 MYCOLOGIA
York) under conditions listed (TABLE I). PCR products were
cleaned with ExoSAP-ITH (USB Corp., Cleveland, Ohio).
Clean PCR products were sequenced at the DNA Sequenc-
ing Facility (Center for Agricultural Biotechnology, Univer-
sity of Maryland, College Park, Maryland) and McLAB DNA
sequencing services (San Francisco, California). Sequences
were assembled and edited with Sequencher 4.9 (Gene
Codes Corp., Madison, Wisconsin). Sequences were depos-
ited in GenBank (S
UPPLEMENTARY TABLE I).
Phylogenetic analyses.—
Two separate phylogenetic analyses
were performed on two separate datasets as described
below. The first dataset contained a reduced number of
isolates of
Cosmospora vilior
and related taxa as well as
species of other cosmospora-like fungi to elucidate their
phylogenetic placement in the Nectriaceae. The second
dataset contained all isolates of
Cosmospora vilior
and
related taxa to determine their relationships.
ITS-LSU, MCM7,
rpb1
,
tef1
and
tub
sequences were
aligned with MAFFT 6 (Katoh 2008) and manually edited
if necessary in Mesquite 2.75 (Maddison and Maddison
2011). Gaps (insertions/deletions) were treated as missing
data. Alignments were deposited in TreeBASE (http://www.
treebase.org; accession number S14038). Maximum likeli-
hood (ML) and Bayesian (BI) analyses were performed on
each of the datasets of individual loci first and then on the
concatenated dataset. CONCATEPILLAR 1.4 (Leigh et al.
2008) was used to determine whether loci could be analyzed
by concatenating the datasets or whether loci should be
analyzed separately. Loci were concatenated if the
P
value
was greater than the default a-level of 0.05, which indicated
that the null hypothesis (i.e. congruence of loci) could not
be rejected.
For both ML and BI analyses, jModeltest (Guindon and
Gascuel 2003, Posada 2008) was used to infer the models of
nucleotide substitution for each locus. Default settings in
jModeltest were used: 11 substitution schemes with equal or
unequal base frequencies (+F) and invariable sites (+I)
and/or rate variation among sites (+G). The base tree for
likelihood calculations was ML optimized. Once likelihood
scores were calculated, the models were selected according
to the Akaike information criterion (AIC).
Maximum likelihood (ML) analyses were performed with
GARLI 2.0 (Genetic algorithm for rapid likelihood infer-
ence; Zwickl 2006) by submitting the job via the GARLI web
service at http://www.molecularevolution.org (Bazinet and
Cummings 2011), which uses a grid computing system
(Cummings and Huskamp 2005) associated with the Lattice
Project (Bazinet and Cummings 2008). Fifty independent
search replicates were performed to find the best tree. The
starting tree was generated with a fast ML stepwise-addition
algorithm. Two thousand bootstrap replicates were used for
bootstrap analysis. Bayesian analyses were performed in
MrBayes 3.2.1 (Ronquist et al. 2012). A majority-rule
consensus tree was generated by running four chains for
10 000 000 Markov chain Monte Carlo generations, sam-
pling trees every 100th generation and discarding the first
25% of the sampled trees as burn-in. Tracer 1.5 (Rambaut
and Drummond 2007) was used to confirm whether the
negative log likelihoods had reached convergence.
RESULTS
Phylogenet ic analyses: phylogenetic placement of C.
vilior and related species within Cosmospora sensu
Rossman.—
The analysis performed in CONCATE-
PILLAR failed to reject the null hypothesis of
congruence among loci (
P
5 0.08). Therefore all loci
were concatenated. The concatenated matrix included
22 ingroup isolates that formed five major groups plus
two outgroup taxa (
Corallomycetella repens
and
Pseudo-
nectria pachysandricola
). It consisted of 3591 base pairs
of which 970 were parsimony informative, 318 were
parsimony uninformative and 1858 were invariable
sites. The topologies of the generated phylogenetic
trees in both ML and BI were congruent. The negative
log likelihoods for the phylogenetic trees were
216460.154 and 216513.115 respectively. The best
tree (ML) is illustrated (F
IG. 1).
Cosmospora vilior
,
C. joca
and related species
formed a highly supported clade (94% BP, 100%
PP). This clade is related to
Dialonectria
,
Cosmospora
sensu stricto and an orphan group that includes
C.
flavoviridis
,
C. obscura
and
C. stegonsporii
. These
clades of cosmospora-like fungi were highly support-
ed as well (.70% BP, .95% PP), but the inner nodes
connecting these clades were poorly supported. Basal
to all of these groups is
Microcera
, another segregate
genus of cosmospora-like fungi.
Phylogenetic analyses: relationship among C. vilior and
related species.—
The null hypothesis of congruence
among loci (
P
5 0.11) was not rejected in CON-
CATERPILLAR, and therefore the five loci were
concatenated to estimate a phylogeny. The concate-
nated matrix included 25 isolates belonging to the
ingroup and two outgroup taxa (
C. repens
and
M.
larvarum
). The concatenated matrix consisted of
3353 bp of which 651 were parsimony informative,
456 were parsimony uninformative and 1785 invari-
able sites. The tree topologies generated with ML and
BI were congruent. The log likelihoods for these two
analyses were 223 024.3191 and 223 055.9286 respec-
tively. The best tree generated with ML is illustrated
(F
IG. 2).
The combined analyses of
Cosmospora vilior
and
related species revealed that there were as many as 16
independently evolving lineages (5 putative species).
Clade I is a complex of species whose hosts are
Eutypella
species. Three species are recognized within
this clade, which include
C. vilior
and two species
described below (
Pseudocosmospora eutypellae
,
P.
rogersonii
). Sister to clade I is
P. eutypae
(described
below), whose host is a species of
Eutypa
Tul. & C.
Tul. Sister to clade II (
P. eutypae
+ clade I) is clade III,
which comprises two monotypic species,
C. joca
and
P. metajoca
(described below).
Cosmospora joca
is
H
ERRERA ET AL.:
P
SEUDOCOSMOSPORA
1289
TABLE I. Genes/loci used in the phylogenetic analyses
Locus ITS LSU Mcm7 Rpb1 Tef1 Tub Combined
Cosmospora
sensu lato
dataset
Nucleotide substitution
models
TIM2+I+2 TrN+I+G TIM2+I+G TrN+I+G TPM3uf+I+G
Included sites 1384 570 692 365 580 3591
Phylogenetically
informative sites
94 201 235 236 204 970
Uninformative
polymorphic sites
77 29 80 42 90 318
Invariable sites 1028 324 282 25 199 1858
Pseudocosmospora
dataset
Nucleotide substitution
models
TIMef+I+G TrN+I+G TIM2+I+G HKY+G TIM3+I+G
Included sites 1323 561 642 280 547 3353
Phylogenetically
informative sites
65 44 211 172 159 651
Uninformative
polymorphic sites
64 166 70 64 92 456
Invariable sites 992 327 253 0 213 1785
Primers used (reference) ITS5, ITS4
(White et al.
1990)
LR5, LROR
(Vilgalys and
Hester 1990)
mcm7-709for,
mcm7-1348rev
(Schmitt et al. 2009)
crpb1a, rpb1c
(Castlebury et al.
2004)
tef1-728, tef1-986
(Carbone and
Kohn 1999)
Btub-TI, Btub-T2
(O’Donnell and
Cigelnik 1997)
PCR protocol: annealing temperature
and cycles
53 C, 1 min, 403 56 C, 50 s, 383 50 C, 2 min, 403 66 C, 55 s, 93
56 C, 55 s, 353
55 C, 30 s, 353
1290 MYCOLOGIA
associated with a species of
Biscogniauxia
Kuntze
(Xylariaceae), while
P. metajoca
is associated with a
species of
Eutypa
. All clades corresponding to
recognized species received maximum BP and PP
support (with one exception).
T
AXONOMY
Pseudocosmospora C. Herrera & P. Chaverri, gen.
nov.
MycoBank MB802432
Type species: Pseudocosmospora eutypellae
C. Herrera
& P. Chaverri
Etymology:
‘‘
Pseudo
’’ from Greek referring to the mor-
phological similarity to both the teleomorphic and anamor-
phic states of
Cosmospora
sensu stricto.
Teleomorph:
Stroma absent. Perithecia superficial or
slightly immersed in fungal host stroma, scattered to
gregarious, subglobose to obpyriform with a blunt
papilla, generally less than 250
mm high, soft-textured,
smooth-walled, scarlet, KOH+ blood red, LA+ yellow,
collapsing laterally when dry, uniloculate. Perithecial
surface cells forming textura angularis. Perithecial
wall generally 20–30
mm thick, of two regions, outer
region of cells forming textura globulsa to t.
angularis; inner region of cells forming textura
prismatica. Asci unitunicate, cylindrical to narrowly
clavate, increasing in size as ascospores mature,
without a conspicuous apical ring, with eight spores
arranged uniseriately. Ascospores ellipsoidal, one-
septate, slightly constricted at septum, yellow-brown,
verrucose, sometimes appearing smooth at maturity.
Anamorph in culture:
After 21 d at room tempera-
ture on PDA, colony surface crustose with no aerial
mycelium or cottony with aerial mycelium, rosy-buff,
pale-luteous, or salmon-pink. Sporulation on SNA
FIG. 1. Phylogenetic placement of
C. vilior
and related species within
Cosmospora
sensu Rossman based on a combined five-
loci (ITS-LSU, MCM7,
rpb1
,
tef1
,
tub
) dataset. Best tree generated with ML analysis (216 460.154). Values at branches indicate
maximum likelihood bootstrap (ML BP)/Bayesian posterior probabilities (BI PP).
HERRERA ET AL.:
P
SEUDOCOSMOSPORA
1291
usually abundant, arising directly from agar surface.
Anamorphic state acremonium-like to verticillium-
like; conidiophores generally simple, unbranched,
sometimes verticillately branched, rarely densely
aggregated. Phialides monophialidic, cylindrical, hy-
aline. Conidia ellipsoidal, ovoid, or reniform, smooth,
sometimes guttulated, non-septate, hyaline.
Habitat:
On stromata of diatrypaceous fungi,
particularly species of
Eutypa
and
Eutypella
, rarely
on species of
Biscogniauxia
.
Distribution:
Asia, Africa, Europe, North America,
Oceania, South America, possibly cosmopolitan.
Notes: Pseudocosmospora
is similar to
Cosmospora
sensu stricto in its cosmospora-like teleomorph and
acremonium-like anamorph, although they differ in
cultural characteristics and host preference.
Pseudo-
cosmospora
is most common on diatrypaceous fungi
except for
P. joca
, which occurs on a
Biscogniauxia
sp.
The latter does not belong among the known hosts of
Cosmospora
sensu stricto, although these attack
xylariaceous fungi as well as polypores. In general,
s
pecies of
Pseudocosmospora
have pinkish colonies,
while species of
Cosmospora
sensu stricto have
olivaceous green colonies on PDA. Phylogenetically
Pseudocosmospora
appears to be closely related to
Dialonectria
. Both occur on diatrypaceous fungi,
although they attack different genera. The genera
also differ in their anamorphic state;
Dialonectria
has
a fusarium-like anamorph.
K
EY TO SPECIES OF
P
SEUDOCOSMOSPORA
1. On
Biscogniauxia
(Xylariaceae) ..........
P. joca
1. On
Eutypa
or
Eutypella
(Diatrypaceae) . ....... 2
FIG. 2. Phylogenetic relationship of
C. vilior
and related species based on a combined five-loci (ITS-LSU, MCM7,
rpb1
,
tef1
,
tub
) dataset. Best tree generated with ML analysis (223 024.3191). Values at branches indicate maximum likelihood bootstrap
(ML BP)/Bayesian posterior probabilities (BI PP).
1292 MYCOLOGIA
2. On
Eutypa
............................. 3
2. On
Eutypella
............................ 4
3. Ascospores smooth, 6.3–8.7
mm long; colony rosy
buff, 6–15 mm diam after 14 d at 25 C on PDA . .
..........................
P. eutypae
3. Ascospores verrucose, 7.7–11.9 mm long; colony
salmon-pink, 24–25 mm diam after 14 d at 25 C on
PDA ..........................
P. metajoca
4. Perithecia with a discoidal apex . . . ........... 5
4. Perithecia with a blunt apex ................ 6
5. Fungal host on
Alnus
sp.; ascospores smooth, 9–
10.4
mm .......................
P. pithoides
5. Fungal host on
Espeletia
sp.; ascospores verrucose,
11–14
mm long ............
P. pseudepisphaeria
6. Conidiophores branching, becoming densely ra-
mulose (fasciculate) on SNA ..........
P. triqua
6. Conidiophores not branching or sparingly
branched on SNA ........................ 7
7. Colonies pale-luteous on PDA; conidia reniform,
with two guttules at opposite ends, 3.4–7.4
mm
long .............................
P. vilior
7. Colonies white to salmon-pink on PDA ........ 8
8. Colonies white on PDA; ascospores smooth, 10–
15
mm long .................
P. metepisphaeria
8. Colonies salmon-pink on PDA ............... 9
9. Ascospores verrucose, 7.1–12.5
mm long; colonies
7.5–20 mm diam after 14 d at 25 C on PDA; conidia
oblong to ellipsoidal, with two guttules at opposite
ends, 3.1–6.2
mm long . ............
P. eutypellae
9. Ascospores smooth, 7.9–12.2 mm long; colonies 18–
27.5 mm diam after 14 d at 25 C on PDA; conidia
oblong to ellipsoidal, without guttules, 2.9–5.5
mm
long ..........................
P. rogersonii
Pseudocosmospora eutypae C. Herrera & P. Cha-
verri, sp. nov. F
IG.3.
MycoBank MB802433
Holotype:
FRANCE, Poitou-Charentes, Saint George
de Rex (Marais Poitevin), on
Eutypa
sp., 26 Apr 2011,
C. Herrera
(C.H. 11-01), BPI 884164, ex-holotype
culture CBS 133961.
Etymology:
In reference to its fungal host,
Eutypa
.
Teleomorph:
Perithecia solitary, superficial, nonstro-
matic, subglobose with a discoidal apex, collapsing
laterally when dry, scarlet, smooth, 171–200 3 150–
183
mm (mean 5 182 3 165; SD 16.1, 16.5; n 5 3).
Asci cylindrical to slightly clavate, with eight spores
arranged uniseriately, 54–66 3 5.5–7
mm (mean 5 59
3 6; SD 5.4, 0.7; n 5 4). Ascospores ellipsoid, equally
two-celled, one-septate, slightly constricted at septum,
smooth, hyaline, 6.3–8.7 3 3.1–4.1
mm (mean 5 7.8 3
3.6; SD 0.7, 0.3; n 5 30).
Anamorph:
Colonies 6–15 mm diam (mean 5 11.4;
SD 4.1; n 5 5) after 14 d. at 25 C on PDA, cottony with
rosy-buff aerial mycelium, reverse concolorous. Spor-
ulation on SNA usually abundant, arising directly
from agar surface. Anamorphic state acremonium-
like; conidiophores generally simple, unbranched.
Phialides monophialidic, cylindrical, collarette not
flared, hyaline, length 36–53
mm (mean 5 41.3; SD
4.1; n 5 13), width at base 1.5–2.0
mm (mean 5 1.8;
SD 0.2; n 5 13), width at tip 1–1.3
mm (mean 5 1.1;
SD 0.1; n 5 13). Conidia oblong, unicellular, smooth,
hyaline, 4.6–6.7 3 1.2–2.1
mm (mean 5 5.7 3 1.7; SD
0.6, 0.2; n 5 30).
Habitat:
On
Eutypa
cf.
lata
(Diatrypaceae) bark.
Distribution:
France, United Kingdom.
Additional isolates examined:
UNITED KINGDOM, on
Crataegus
sp., 1958,
S. Francis
, culture IMI 73016.
Notes: Pseudocosmospora eutypae
occurs on
Eutypa
cf.
lata
and has small and smooth ascospores.
Pseudocosmospora metajoca
is the only other species
of
Pseudocosmospora
on
Eutypa
, but it has longer and
verrucose ascospores.
Pseudocosmospora eutypellae C. Herrera & P.
Chaverri, sp. nov. F
IG.4.
MycoBank MB802434
Holotype:
USA, Maryland, Beltsville, on
Eutypella
sp., on dead twigs of unidentified tree, 7 Oct 2008,
Y
.
Hirooka
(Y.H. 08-17), BPI 884165, ex-holotype culture
CBS 133966 5 A.R. 4562.
Etymology:
In reference to its fungal host,
Eutypella
.
Teleomorph:
Perithecia gregarious, slightly im-
mersed in host stromata, subglobose with a blunt
apex to obpyriform, collapsing laterally, scarlet,
smooth, 143–303 3 108–205
mm (mean 5 202 3
150; SD 39, 22.7; n 5 22). Asci cylindrical to slightly
clavate, eight-spored, uniseriately arranged, 63–78.6
3 5.9–7.9
mm (mean 5 71.6 3 6.7; SD 4.5, 0.6; n 5
27). Ascospores ellipsoid to fusiform, equally two-
celled, slightly verrucose, yellow-brown, 7.1–12.5 3
3.6–5.6
mm (mean 5 9.9 3 4.5; SD 0.9, 0.4; n 5 136).
Anamorph:
Colonies 7.5–20 mm diam (mean 5
12.6, SD 3.4, n 5 18) after 14 d. at 25 C on PDA,
sometimes crustose, with or without aerial mycelium,
buff, rosy-buff or salmon-pink, reverse concolorous.
Sporulation on SNA usually abundant, arising directly
from agar surface, sometimes from lateral pegs.
Anamorphic state acremonium-like to verticillium-
like; conidiophores simple, unbranched, or
branched, becoming densely branched. Phialides
monophialidic, cylindrical, collarette not flared,
hyaline, length (3.8–)7.8–15.1(–18.5)
mm (mean 5
10.2; SD 3.0; n 5 29), width at base 0.9–2.3
mm (mean
5 1.3; SD 0.3; n 5 29), width at tip 0.7–1.2
mm (mean
5 0.9; SD 0.14; n 5 29). Conidia oblong to ellipsoidal,
unicellular, with two guttules at opposite ends,
smooth, hyaline, 3.1–6.2 3 1.0–2.4
mm (mean 5 4.2
3 1.5; SD 0.6, 0.2; n 5 1 80).
Habitat:
On
Eutypella
sp. (Diatrypaceae) bark.
H
ERRERA ET AL.:
P
SEUDOCOSMOSPORA
1293
FIG.3.
Pseudocosmospora eutypae
. A. Perithecia on natural substrata. Bar 5 200 mm. B. Perithecium in 3% KOH. Bar 5
100
mm. C. Median section of perithecium. Bar 5 100 mm. D. Perithecial surface cells. Bar 5 100 mm. E. Asci. Bar 5 10 mm. F.
Ascospore. Bar 5 10
mm. G. Cultures after 3 wk at 25 C on PDA. Bar 5 10 mm. H. Phialide. Bar 5 10 mm. I. Conidia. Bar 5
10
mm.
1294 MYCOLOGIA
FIG.4.
Pseudocosmospora eutypellae
. A, B. Perithecia on natural substrata. A. Bar 5 600 mm; B. bar 5 200 mm. C. Perithecium
in 3% KOH. Bar 5 100
mm. D. Median section of perithecium. Bar 5 100 mm. E. Perithecial surface cells. Bar 5 100 mm. F. Asci.
Bar 5 10
mm. G. Ascopore. Bar 5 10 mm. H, I. Cultures after 3 wk at 25 C on PDA. Bars 5 10 mm. J, K. Phialides. Bars 5 10 mm.
L. Lateral phialidic pegs. Bar 5 10
mm. M. Conidia. Bar 5 10 mm.
HERRERA ET AL.:
P
SEUDOCOSMOSPORA
1295
Distribution:
France and U.S.A.
Additional specimens and isolates examined:
FRANCE,
Oloron, Foreˆt de Bugangue, on
Eutypella
sp., on bark of
Robinia pseudoacacia
(?), 17 May 1993,
F. Candoussau &
J.D. Rogers
(F. 262), BPI 802567, culture CBS 128986 5
G.J.S. 93-15; USA, Kentucky, Clermont, Bernheim Arbore-
tum and Research Forest, on
Eutypella
sp., on dead branch
of unidentified tree, 27 Jun 2010,
Y. Hirooka
, BPI 884169,
culture CBS 133977 5 G.J.S. 10-248; Maryland, Frederick
County, Cunningham Falls State Park, on
Eutypella
sp., on
Rhus typhina
, 26 Aug 2007,
L. Vasilyeva
, BPI 878454,
culture CBS 129430 5 A.R. 4453; Pennsylvania, Greensburg,
on
Eutypella
sp., Aug 2008,
J. Plitschke
, culture CBS 133965
5 A.R. 4527; West Virginia, Grafton, on
Eutypella
sp., on
bark of unidentified tree, 26 Jun 2010,
Y. Hirooka
, BPI
884168, culture CBS 133960 5 C.H. 10-02.
Notes: Pseudocosmospora eutypellae
is most closely
related and similar to
P. rogersonii
but can be
distinguished from the latter by the ornamentation of
its ascospores.
Pseudocosmospora eutypellae
has verrucose
ascospores, while
P. rogersonii
has smooth ascospores.
Pseudocosmospora joca (Samuels) C. Herrera & P.
Chaverri, comb. nov. F
IG.5.
Mycobank MB802435
Basionym: Nectria joca
Samuels, Mycol. Pap. 164:21. 1991.
;
Cosmospora joca
(Samuels) Rossman & Samuels, Stud.
Mycol. 42:122. 1999.
Teleomorph:
Perithecia gregarious, superficial, non-
stromatic, subglobose with a minute papilla, collaps-
ing laterally, scarlet at first, becoming blood red,
darker at apex, smooth, 375–384 3 317–349
mm
(mean 5 380.2 3 336.3; SD 4.9, 16.9; n 5 3). Asci
cylindrical to clavate, eight-spored, uniseriately ar-
ranged, 90.7–112.6 3 8.8–11.1
mm (mean 5 102.2 3
9.8; SD 8.5, 0.7; n 5 9). Ascospores ellipsoid, equally
two-celled, one-septate, constricted at septum, verru-
cose, yellow-brown, 10.9–14 3 6.4–7.7
mm (mean 5
12.8 3 7; SD 0.8, 0.3; n 5 30).
Anamorph:
Colonies 4 mm diam (n 5 3) after 14 d.
at 25 C on PDA, crustose, salmon-pink to orange
colony, reverse concolorous. Rarely sporulating on
SNA. Anamorphic state acremonium-like; conidio-
phores generally simple, unbranched. Phialides
monophialidic, cylindrical, hyaline, length 14.3–
24.2
mm (mean 5 18.8; SD 5.0; n 5 3), width at base
1.6–2.4
mm (mean 5 2.0; SD 0.4; n 5 3), width at tip
1.1 (n 5 3). Conidia oblong, unicellular, smooth,
hyaline, 3.0–5.5 3 1.3–2.1
mm (mean 5 4.1 3 1.6; SD
0.6, 0.2; n 5 30).
Habitat:
On
Biscogniauxia
sp. bark.
Distribution:
Argentina and Brazil.
Holotype:
BRAZIL, Amazonas, Pico Rondon, Km
211 on Perimetral Norte, ca. 3 h walk from FUNAI
post toward summit, 01u329N, 02u489W, on
Biscog-
niauxia
sp., 25 Mar. 1984,
G.J. Samuels
(1094),
Pipoly
& Guedes
, INPA (not seen), ISOTYPES BPI 802606,
NY 00671973 (not seen).
Epitype designated herein:
ARGENTINA, ´o Negro
Province, San Carlos de Bariloche, Luma forest, on
Biscogniauxia
sp., on rotted wood, 15 Apr 2011,
A.
Romero
, BPI 884175, ex-epitype culture CBS 133967 5
A.R. 4779.
Notes:
The application of the name is restricted
here to species of
Pseudocosmospora
on
Biscogniauxia
.
The isotype and the designated epitype both occur on
species of
Biscogniauxia
. The colony and the ana-
morph are similar to the morphology described in the
original description, although the perithecia and
ascospores of the epitype are larger than those
reported in the literature.
Pseudocosmospora metajoca C. Herrera & P. Cha-
verri, sp. nov. F
IG.6.
MycoBank MB802436
Holotype:
NEW ZEALAND, North Island, Mount
Williams, on
Eutypa
sp., on dead woody branch of
Beilschmiedia tawa
, 7 Mar 2009,
A.Y. Rossman & P.
Chaverri
(P.C. 952), BPI 879088, ex-holotype culture
CBS 133968 5 A.R. 4576.
Etymology:
‘‘
Meta
’’ from Greek meaning adjacent and
‘‘
joca
’ in reference to the fact that it originally was classified
as
C. joca
, and later found to be phylogenetically close to
C.
joca
.
Teleomorph:
Solitary or gregarious, superficial, non-
stromatic, subglobose with a discoidal apex, some
collapsing laterally, scarlet, smooth, 222–251 3 204–
213
mm (mean 5 236 3 208; n 5 2). Asci clavate,
eight-spored, uniseriately arranged, 62.2–69.2 3 5.9–
7.2
mm (mean 5 65.5 3 6.4; SD 2.5, 0.5; n 5 5).
Ascospores ellipsoid, equally two-celled, one-septate,
slightly constricted at septum, slightly verrucose,
yellow-brown, 7.7–11.9 3 3.3–5.3
mm (mean 5 8.9 3
4.3; SD 0.9, 0.5; n 5 29).
Anamorph:
Colonies 24–25 mm diam (mean 5
24.5; SD 0.5; n 5 3) after 14 d. at 25 C on PDA,
slightly cottony, pale salmon-pink, reverse concolor-
ous. Sporulation on SNA usually abundant, arising
directly from agar surface. Anamorphic state acremo-
nium-like; conidiophores generally simple, un-
branched. Phialides monophialidic, cylindrical, col-
larette not flared, hyaline, length 26–49
mm (mean 5
37.3; SD 5.7; n 5 9), width at base 2.1–2.8
mm (mean
5 2.4; SD 0.2; n 5 9), width at tip 1–1.3
mm (mean 5
1.2; SD 0.1; n 5 9). Conidia oblong to ellipsoidal,
unicellular, guttulated, smooth, hyaline, 3.8–6.1 3
1.6–3.1
mm (mean 5 4.8 3 2.1; SD 0.6, 0.3; n 5 30).
Habitat:
On
Eutypa
sp. (Diatrypaceae) on dead
branch of
Beilschmiedia tawa
.
Distribution:
New Zealand.
1296 M
YCOLOGIA
Notes: Pseudocos mospora metajoca
originally was
identified as
C. joca
based on its occurrence on what
was thought to be a stroma of a
Biscogniauxia
species
and its salmon-pink culture on PDA. On close
examination of the specimen, the host was found to
be a species of
Eutypa
. The colony of
P. metajoca
grows faster than of
P. joca
. In addition,
P. metajoca
has much smaller perithecia and ascospores com-
FIG.5.
Pseudocosmospora joca
. A, B. Perithecia on natural substrata. A. Bar 5 2 mm; B bar 5 200 mm. C. Asci. Bar 5 10 mm.
D. Ascospores. Bar 5 10
mm. E. Cultures after 3 wk at 25 C on PDA. Bar 5 10 mm. F. Phialide. Bar 5 10 mm. G. Conidia. Bar 5
10
mm.
HERRERA ET AL.:
P
SEUDOCOSMOSPORA
1297
pared to
P. joca
.
Pseudocosmospora metajoca
differs
from
P. eutypae
, the other
Pseudocosmospora
on
Eutypa
, by having verrucose ascospores.
Pseudocosmospora metepisphaeria (Samuels) C. Her-
rera & P. Chaverri, comb. nov.
MycoBank MB802437
Basionym: Nectria metepisphaeria
Samuels, Mycol. Pap.
164:29. 1991.
;
Cosmospora metepisphaeria
(Samuels) Rossman &
Samuels, Stud. Mycol. 42:123. 1999.
Anamorph:
Acremonium-like
Habitat:
On
Eutypella
sp. (Diatrypaceae) on un-
identified bark.
FIG.6.
Pseudocosmospora metajoca
. A, B. Perithecia on natural substrata. A. Bar 5 2 mm; B. bar 5 200 mm. C. Perithecium in
3% KOH. Bar 5 100
mm. D. Median section of perithecium. Bar 5 100 mm. E. Asci. Bar 5 10 mm. F. Ascospores. Bar 5 10 mm.
G. Cultures after 3 wk at 25 C on PDA. Bar 5 10 mm. H, I. Phialides. Bars 5 10
mm. J. Conidia. Bar 5 10 mm.
1298 MYCOLOGIA
Distribution:
Venezuela (known only from the type
collection)
Holotype:
VENEZUELA, Dist. Federale, vic. Ma-
carao, on
Eutypella
sp., on unidentified bark, 21 Jun
1971,
K.P. Dumont
(VE 335),
J.H. Haines
,
G. Morillo
& E. Moreno
, VEN (not seen), ISOTYPE NY.
Notes:
The isotype specimen was studied and
determined to occur on a
Eutypella
sp. Based on this
host, it can be predicted that
C. metepisphaeria
would
fall within the
Pseudocosmospora
clade. In addition to
the host, the reported acremonium-like anamorphic
state supports the placement of this species in
Pseudocosmospora
. Unique to this species is its smooth
ascospores, (10–)11–14(–15)
mm long and the white,
crustose colony on PDA, reverse brown (Samuels et al.
1991). A culture no longer exists.
Pseudocosmospora pithoides (Ellis & Everh.) C.
Herrera & P. Chaverri, comb. nov.
MycoBank MB802438
Basionym: Nectria pithoides
Ellis & Everh., Proc. Acad. Nat.
Sci. Philad. 43:247 (1891).
Anamorph:
Unknown
Habitat:
On an
Eutypella
sp. (Diatrypaceae) bark of
dead alder.
Distribution:
British Columbia (known only from
the type collection).
Holotype:
CANADA, British Columbia, bark of dead
alder, May 1889,
J. Macoun
(122), NY 00927939.
Notes:
The holotype specimen of
Nectria pithoides
was examined and determined to agree with the
concept of
Pseudocosmospora
in regard to the host,
which appears to be a
Eutypella
species. The
perithecia have a prominent discoidal apex, which
according to the description gives an impression of
being barrel-shaped (
pithos
from Greek 5 barrel). No
asci were observed. The ascospores are ellipsoidal,
one-septate, slightly constricted at the septum,
smooth, 9–10.4 3 4.1–4.5
mm (mean 5 9.7 3 4.4;
SD 0.5, 0.1; n 5 8).
Pseudocosmospora pseudepisphaeria (Samuels) C.
Herrera & P. Chaverri, comb. nov.
MycoBank MB802439
Basionym: Nectria pseudepisphaeria
Samuels, Mycol. Pap.
164:34. 1991.
; C
osmospora pseudepisphaeria
(Samuels) Rossman &
Samuels, Stud. Mycol. 42:124. 1999.
Anamorph:
Acremonium-like.
Habitat:
On
Eutypella
sp. (Diatrypaceae) on branch
of
Espeletia
sp.
Distribution:
Venezuela (known only from the type
collection).
Holotype:
VENEZUELA, Merida, Parque Nacional
Sierra Nevada, near Apartaderos, E. of Laguna
Mucubaji, Laguna Negra, on
Eutypella
sp., on
Espeletia
sp., 18 Jul 1971,
K.P. Dumont
(VE 2277),
J.H. Haines
,
G.J. Samuels & A. Revas
, NY 01013169.
Notes:
Based on our examination of the holotype
specimen, the fungal host of
C. pseudepisphaeria
is a
Eutypella
sp. The fungal host and the reported
acremonium-like anamorphic state support the place-
ment of
C. pseudepisphaeria
in the genus
Pseudocos-
mospora
. Unique to this species are the discoidal
perithecial apices, its verrucose, (11–)11.2–13(–14)
mm long ascospores, and its white to pale salmon-
colored colony (Samuels et al. 1991). A culture no
longer exists.
Pseudocosmospora rogersonii C . Herrera & P.
Chaverri, sp. nov. F
IG.7.
MycoBank MB802440
Holotype:
USA, New York, Dutchess County, Pawl-
ing, Pawling Nature Reserve, on
Eutypella
sp., 6–8 Oct.
1990,
G.J. Samuels & C.T. Rogerson
, BPI 1107121, ex-
holotype culture CBS 133981 5 G.J.S. 90-56.
Etymology:
In honor of Clark T. Rogerson for his work on
the Hypocreales that has guided all of us.
Teleomorph:
Perithecia gregarious, slightly im-
mersed in host stromata, subglobose with a blunt
papilla, collapsing laterally, scarlet, smooth, 163–245
3 131–180
mm (mean 5 193 3 152; SD 37, 21; n 5 7).
Asci broadly cylindrical to narrowly clavate, eight-
spored, uniseriately arranged, 54–69 3 5.7–8.4
mm
(mean 5 63 3 6.7; SD 4.9, 0.7; n 5 12). Ascospores
ellipsoid, equally two-celled, one-septate, slightly
constricted at septum, smooth, yellow-brown, 7.9–
12.2 3 3.3–4.9
mm (mean 5 9.6 3 4.1; SD 0.9, 0.3; n 5
86).
Anamorph:
Colonies 18–27.5 mm diam (mean 5
22.4; SD 3.4; n 5 8) after 14 d at 25 C on PDA,
crustose, rosy-buff to salmon-pink, reverse concolor-
ous. Sporulation on SNA usually abundant, arising
directly from agar surface. Anamorphic state acremo-
nium-like; conidiophores generally simple, un-
branched. Phialide cylindrical, smooth, straight,
collarette not flared, hyaline, length 6.8–29.4
mm
(mean 5 12.3; SD 5; n 5 30), width at base 1.0–2.3
mm
(mean 5 1.5; SD 0.3; n 5 30), width at tip 0.7–1.2
mm
(mean 5 0.9; SD 0.13; n 5 30). Conidia oblong to
ellipsoidal, unicellular, smooth, hyaline, 2.9–5.5 3
1.1–2.6
mm (mean 5 3.8 3 1.6; SD 0.6, 0.3; n 5 89).
Habitat:
On
Eutypella
sp. (Diatrypaceae) bark.
Distribution:
USA.
Additional specimens and isolates examined:
USA, New
York, Dutchess County, Pawling, Pawling Nature Reserve, on
Eutypella
sp., 6–8 Oct 1990,
G.J. Samuels & C.T. Rogerson
,
BPI 1107120; New York, Huguenot, YMCA Greenkill Retreat
Center, on
Eutypella
sp., 26 Sep 2009,
C. Herrera
(C.H. 09-
02), BPI 884167, culture 5 G.J.S. 09-1384; New York,
HERRERA ET AL.:
P
SEUDOCOSMOSPORA
1299
Painted Post, Watson Homestead Conference and Retreat
Center, on
Eutypella
sp., on dead branch of
Fagus grand-
ifolia
, 17 Sep 2010,
C. Herrera
(C.H. 10–11), BPI 884166,
culture CBS 133978 5 G.J.S. 10-296; New York, Painted Post,
Watson Homestead Conference and Retreat Center, on
Eutypella
sp., on dead branch of
Fagus grandifolia
, 17 Sep
2010,
C. Herrera
(C.H. 10-12), BPI 884170, culture CBS
133979 5 G.J.S. 10-297.
Notes: Pseudocosmospora rogersonii
is closely related
to
P. eutypellae
but differs conspicuously in the
ornamentation of its ascospores.
Pseudoscosmospora
rogersonii
has smooth ascospores in contrast to
P.
eutypellae
, which has verrucose ascospores.
Pseudocosmospora triqua (Samuels) C. Herrera & P.
Chaverri, comb. nov.
MycoBank MB802441
Basionym: Nectria triqua
Samuels, Mycol. Pap. 164:40.
1991.
FIG.7.
Pseudocosmospora rogersonii
. A, B. Perithecia on natural substrata. A. Bar 5 2 mm; bar 5 200 mm. C. Perithecium in
3% KOH. Bar 5 100
mm. D. Perithecial surface cells. Bar 5 100 mm. E. Ascus. Bar 5 10 mm. F. Ascospores. Bar 5 10 mm. G, H.
Cultures after 3 wk at 25 C on PDA. Bar 5 10 mm. I. Phialide. Bar 5 10
mm. J. Conidia. Bar 5 10 mm. Bars: A 5 2 mm; B 5
200
mm; C, D 5 100 mm; E,F, I, J 5 10 mm; G, H 5 10 mm.
1300 MYCOLOGIA
;
Cosmospora triqua
(Samuels) Rossman & Samuels,
Stud. Mycol. 42:125. 1999.
Anamorph:
Acremonium-like.
Habitat:
On
Eutypella
sp. (Diatrypaceae) on un-
identified bark.
Distribution:
French Guiana (known only from the
type collection).
Holotype of
Nectria triqua: FRENCH GUIANA,
Upper Marouini River, vic. roche Koutou, 02u559N,
54u049W, 400 m., on
Eutypella
sp., on unidentified
bark, 17 Aug 1987,
G.J. Samuels
(5818),
J.-J. de
Granville
,
L. Allorge
,
W. Hahn
,
M. Hoff
, NY 01013269.
Notes:
Examination of the holotype revealed that
the host is a
Eutypella
sp., which suggests that
C.
triqua
should be placed in the genus
Pseudocosmos-
pora
. In addition, the reported anamorphic state is
similar to that of
P. vilior
and
P. eutypellae
in having
branching conidiophores branch that terminate with
multiple phialides. Cultural morphology in PDA was
not reported in the description of
Nectria triqua
(Samuels et al. 1991). The culture no longer exists.
The ascospores are verrucose and (6.8–)7.8–9.7(–
10.5)
mm long.
Pseudocosmospora vilior (Starba¨ck) C. Herrera & P.
Chaverri, comb. nov. F
IG.8.
MycoBank MB802442
Basionym: Nectria vilior
Starba¨ck, Bih. Kongl. Svenska Vet.-
Acad. Handl. 25:28. 1899.
;
Cosmospora vilior
(Starba¨ck) Rossman & Samuels, Stud.
Mycol. 42:126. 1999.
Teleomorph:
Perithecia gregarious, slightly im-
mersed in host stromata, subglobose with blunt apex,
collapsing laterally, scarlet, smooth, 195–224 3 136–
183
mm (mean 5 213 3 164; SD 9.1, 14.7; n 5 10).
Asci cylindrical to clavate, eight-spored, uniseriately
arranged, 59–81 3 5.3–11.0
mm (mean 5 69 3 7.6; SD
6.2, 1.7; n 5 18). Ascospores ellipsoid, equally two-
celled, one-septate, slightly constricted at septum,
slightly verrucose, yellow-brown, 8.3–13.0 3 4.1–
6.4
mm (mean 5 10.2 3 5.2; SD 1.0, 0.5; n 5 90).
Anamorph:
Colonies 23–65 mm diam (mean 5 49;
SD 16.9; n 5 8) after 21 d at 25 C on PDA, cottony
with pale luteous aerial mycelium, reverse concolor-
ous. Sporulation on SNA usually abundant, arising
directly from agar surface; acremonium-like to verti-
cillium-like; conidiophores simple and unbranched at
first, becoming densely branched. Phialides cylindri-
cal, smooth, straight, collarette not flared, hyaline,
length 5.9–19.5
mm (mean 5 12.9; SD 3.4; n 5 9),
width at base 1.1–1.8
mm (mean 5 1.5; SD 0.2; n 5
29), width at tip 0.7–1.3 mm (mean 5 1.0; SD 0.2; n 5
29). Conidia reniform, unicellular, smooth, with two
guttules at opposite ends, hyaline, 3.4–7.4 3 1.1–
2.3
mm (mean 5 4.8 3 1.7; SD 0.7, 0.3; n 5 90).
Habitat:
On
Eutypella
sp. (Diatrypaceae) bark.
Distribution:
Argentina and Brazil.
Holotype:
BRAZIL, Rio Grande do Sul, Santo
Angelo pr. Cachoaira, on
Eutypella
sp., 12 Jan 1893,
Gustav Malme
(114), S F46424.
Epitype designated herein:
ARGENTINA, Misiones
Province, Iguazu´ Biological Station, on
Eutypella
sp.,
25 Apr 2011,
A.Y. Rossman
,
C. Salgado
,
A. Romero
,
R.
Sanchez
, BPI 884176, ex-epitype culture CBS 133971
5 A.R. 4810.
Additional specimens and isolates examined:
ARGENTINA,
Tucuman Province, Tucuman, on
Eutypella
sp., on standing
dead branch of
Piper tucumanum
, 19 Apr 2011,
A. Romero
,
BPI 884174, culture CBS 133970 5 A.R. 4771; BRAZIL,
Bahia, Igrapiu´na, on
Eutypella
sp., 12 Aug 2010,
P. Chaverri
(P.C. 1246),
O. Liparini Pereira
,
D. Pinho
,
A. Luiz Firmino
,
BPI 884172, culture CBS 133963.
Notes:
An epitype was needed to establish an
anamorph for
P. vilior
and to determine its phyloge-
netic placement. The epitype was selected based on
the relatively proximity to the collecting site of the
holotype. The application of the name is restricted to
species of
Pseudocosmospora
on
Eutypella
from South
America that have pale-luteous colonies on PDA.
However, it is recognized here that
P. vilior
consists of
a species complex.
D
ISCUSSION
Genus concept.—
The generic concept
Cosmospora
sensu stricto is based on its type
Cosmospora coccinea
Rabenh., which has
Verticillium olivaceum
W. Gams as
its anamorph. Although the anamorph bears the
name
Verticillium
Nees, the anamorphic state is
acremonium-like (single phialide, unbranched) to
verticillium-like (branching into multiple phialides).
Accepted species in
Cosmospora
sensu stricto have an
acremonium-like anamorph, and it is the character
that circumscribes the genus (Gra¨fenhan et al. 2011).
Conidiophore branching is not unique to the
anamorph of
C. coccinea
because this also is observed
in some anamorphs in the
Cosmospora viliuscula
species complex.
Pseudocosmospora
(described above) is recognized
as a new genus based on the one-to-one genus
concept suggested by Rossman (1993) to accommo-
date
C. vilior
and related species. The one-to-one
genus concept has been used extensively in the
Ascomycota to delimit genera (e.g. Gra¨fenhan et al.
2011; Luo and Zhang 2010, 2012). Briefly, this genus
concept suggests that a genus should be circum-
scribed based on the correlation of its teleomorph to
its unique anamorph state and vice versa. The groups
circumscribed based on this concept are monophy-
letic and often supported by ecological traits (e.g.
H
ERRERA ET AL.:
P
SEUDOCOSMOSPORA
1301
Gra¨fenhan et al. 2011; Luo and Zhang 2010, 2012). In
Gra¨fenhan et al. (2011), the reported hosts for
members of
Cosmospora
sensu stricto were basidiomy-
cetes (e.g.
Fomitopsis
P. Karst.,
Inonotus
P. Karst. and
Stereum
Hill ex Pers.) and xylariaceous fungi (e.g.
Hypoxylon
Bull.).
Microcera
and
Dialonectria
species
have fusarium-like anamorphs,
Microcera
species are
parasites of scale insects and the lectotype species of
Dialonectria
,
D. episphaeria
, is reported on
Diatrype
stigma
(Hoffm.) Fr. (Diatrypaceae; Booth 1959). The
host of
Dialonectria ullevolea
Seifert & Gra¨fenhan has
not been identified, but it is predicted here that the
host will be a diatrypaceous fungus. The orphan clade
consisting of
C. flavoviridis
(Fuckel) Rossman &
Samuels,
C. stegonsporii
Rossman, Farr & Akulov and
C. obscura
Lowen has not been taxonomically revised
FIG.8.
Pseudocosmospora vilior
. A, B. Perithecia on natural substrata. A. Bar 5 2 mm; B. bar 5 200 mm. C. Perithecium in
3% KOH. Bar 5 100
mm. D. Asci. Bar 5 10 mm. E. Ascospore. Bar 5 10 mm. (F) Cultures after 3 wk at 25 C on PDA. Bar 5
10 mm. G. Phialides. Bar 5 10
mm. H. Conidia. Bar 5 10 mm.
1302 MYCOLOGIA
and might require generic recognition. Species in this
clade have a fusarium-like anamorphs, but little is
known about their fungal hosts. Only the host of
C.
stegonsporii
,
Stegonsporium pyriforme
(Hoffm. : Fr.)
Corda (Diaporthales, Sordariomycetes), has been
identified to species. It is possible that all fungal
hosts of species in this clade are members of the
Diaporthales.
The one-to-one genus concept is ideal for the
circumscription of genera in the Ascomycota because
it forces the study of the holomorph and not only the
teleomorph or anamorph. Such view is crucial in
shifting to one name (Norvell 2011). Discarding
information of either the teleomorph or anamorph
to favor one generic hypothesis over the other may
result in para- or polyphyletic groups. For example, a
weak case could be made to group
Cosmospora
,
Dialonectria
,
Pseudocosmospora
and the orphan clade
that consists of
C. flavoviridis
,
C. stegonsporii
and
C.
obscura
into one genus because they have a cosmos-
pora-like teleomorph and occur generally on Sordar-
iomycetes. However, when the anamorphs are super-
imposed on the phylogeny, a paraphyletic group is
formed with two groups having acremonium-like
anamorphs and the remaining two groups fusarium-
like anamorphs. It suggests that the teleomorph state
is probably a symplesiomorphic character (or ances-
tral), while the anamorph represents a synapo-
morphic character (derived). Moreover, segregation
of the discussed genera is supported by specialization
to different host taxa.
The cosmospora-like teleomorphic state of
Pseudo-
cosmospora
was correlated here to an acremonium-like
anamorph. Our phylogeny (F
IG. 2) demonstrates that
Pseudocosmospora
(BP 100%, PP 100%) is not conge-
neric with
Cosmospora
s.str., the only other group of
cosmospora-like fungi with an acremonium-like ana-
morph (Hirooka et al. 2010, Gra¨fenhan et al. 2011).
The two groups differ primarily in their cultural
characteristics. In general,
Pseudocosmosp
ora produc-
es pinkish colonies while
Cosmospora
sensu stricto
produces olivaceous-green colonies on PDA.
Members of each genus considered in this study
occur only on a particular group of host fungi.
Pseudocosmospora
is reported here to occur primarily
on
Eutypa
and
Eutypella
species (Diatrypaceae) with
the exeption of
Cosmospora joca
(Samuels) Rossman
& Samuels, whose host is a species of
Biscogniauxia
.
The genus
Dialonectria
also occurs on diatrypaceous
fungi but has a fusarium-like anamorphic state as do
species in the genus
Microcera
that occur primarily on
insects.
Species concept.—
Genealogical Concordance Phyloge-
netic Species Recognition was used to delimit species
boundaries (GCPSR; Taylor et al. 2000). According to
this operational species concept, putative species are
clades that are concordant across all single gene trees.
The morphological species recognition also was used
to support the species inferences made when applying
GCPSR. Hence, inferred species may be associated
with unique morphological features that set them
apart from other closely related species.
Another species concept that could be useful in
determining additional characters to delimit species
is the ecological species concept. Ecological niches or
adaptive zones can be used to delimit species
(reviewed in de Queiroz 2007), according to this
species concept. Host, an ecological niche, could be a
character specific to a particular
Pseudocosmospora
species. However, host identification of
Eutypa
and
Eutypella
to species was not possible based on
morphology alone. Identification of the fungal host
based on DNA sequences would resolve this problem.
Moreover, analyzing DNA sequences of cosmospora-
like fungi and their associated fungal hosts would
allow testing the hypothesis of cospeciation. Evidence
for cospeciation would provide independent evidence
for the delimitation of species in this genus.
A problem of GCPSR is that it requires multiple
individuals per species. By definition a clade is formed
by a minimum of two individuals per species
(reviewed in Vinuesa 2010). In the case of this paper,
only a single collection was made for many of the
lineages, and it left us with a dilemma on how to deal
with the many singletons present in the phylogeny
(F
IG. 2). It was decided to use the rule of rarity
(reviewed in Lim et al. 2012) to recognize a singleton,
Pseudocosmospora metajoca
(described below), as a
species. This species is morphologically and ecologi-
cally distinct from species recognized with GCPSR
and other singletons.
Pseudocosmospora metajoca
occurs on an
Eutypa
sp. on
Beilschmiedia tawa
(A.Cunn.) Kirk (Lauraceae), which is a broadleaf
tree native to New Zealand and has verrucose
ascospores. Also supporting the view that
P. metajoca
is a distinct species is the relatively long branch
length, which indicates that there have been multiple
substitutions per site since its segregation.
Cosmospora vilior
represents a case where morphol-
ogy is insufficient to distinguish closely related species.
This species is characterized by its relatively fast
growing, pale luteous colony on PDA. However, the
clade probably represents a species complex given the
highly supported subclade that consists of the strains
AR 4771 and PC 1246. The species complex may
consist of up to three species, but the selected epitype
strain, AR 4810, is considered closer to the true
C. vilior
based on geographical proximity to the original
collecting site of the type specimen and its host.
H
ERRERA ET AL.:
P
SEUDOCOSMOSPORA
1303
Phylogenetic placement of Cosmospora joca.—
The
phylogenetic placement of
Cosmospora joca
, the host
of which is a
Biscogniauxia
sp., is puzzling considering
that other members of
Pseudocosmospora
have an
Eutypa
or
Eutypella
species (Diatrypaceae) as their host. Two
potential explanations for this observation are (i) that
the
Biscogniauxia
sp. represents the ancestral host for
Psedocosmospora
species or (ii) that the
Biscogniauxia
host of
C. joca
represents an independent host shift.
Given that
Pseudocosmospora
species have diatrypaceous
and xylariaceous hosts and assuming the first view,
Pseudocosmospora
could represent a link in the diver-
gence from
Cosmospora
sensu stricto to
Dialonectria
(or
vice versa). Phylogenies of the fungal hosts have placed
the Diatrypaceae as a sister clade to Xylariaceae (Moster
et al. 2004, Tang et al. 2009), and these cosmospora-like
fungi could have tracked their hosts faithfully as they
diverged. Host specificity is not uncommon in the
Hypocreales (e.g. species of
Cordyceps
sensu lato are
known to be host specific to insect species; Sung et al.
2007). Co-evolution/cospeciation analyses are needed
to test these hypotheses.
A
CKNOWLEDGMENTS
We gratefully acknowledge the assistance of the curators
and their staff of the herbaria from which specimens were
generously loaned. These herbaria include U.S. National
Fungus Collection (BPI), William and Lynda Steere
Herbarium, New York Botanical Garden (NY) and Herbar-
ium of the Botany Department, Swedish Museum of
National History (S). We also are grateful for the help of
Yuuri Hirooka (Forestry and Forest Products Research
Institute, Japan), Peter Johnston (Landcare Research, New
Zealand), Carlos Mendez (University of Costa Rica), John
Plitschke (Pennsylvania), Andrea Romero and Romina
Sanchez (Departamento de Biodiversidad y Biologia Exper-
imental, Universidad de Buenos Aires, Argentina) and
Catalina Salgado (PSLA, UMD, USA) for contributing in
the organization and collection of various specimens on
collecting trips. The first author especially thanks Chun-
Juan Wang (State University of New York, College of
Environmental Science and Forestry) for mentoring the
author as an undergraduate and guiding the author into
the field of mycology. The first author also thanks current
and former colleagues at USDA-ARS, SMML (USA) and
PSLA, University of Maryland (USA) laboratories, for their
moral support. This study was financially supported by the
United States National Science Foundation (NSF) PEET
grant DEB-0731510 Monographic Studies in the Nectria-
ceae, Hypocreales:
Nectria
,
Cosmospora
, and
Neonectria
to P.
Chaverri, A.Y. Rossman and G.J. Samuels.
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Supplementary resource (1)

... Seven species are currently known in the genus [1,2,5]. The genus Pseudocosmospora, typified by P. eutypellae C.S. Herrera & P. Chaverri, was established by Herrera et al. [6] to accommodate Cosmospora vilior (Starbäck) Rossman & Samuels and related species that are usually fungicolous. Sixteen species are recognized [6][7][8][9]. ...
... The genus Pseudocosmospora, typified by P. eutypellae C.S. Herrera & P. Chaverri, was established by Herrera et al. [6] to accommodate Cosmospora vilior (Starbäck) Rossman & Samuels and related species that are usually fungicolous. Sixteen species are recognized [6][7][8][9]. The genus Thelonectria, typified by T. discophora (Mont.) ...
... The accumulated morpho-logical and phylogenetic information indicated that the genus was not monophyletic [41,42]. Herrera et al. [6] established Pseudocosmospora to accommodate ten cosmospora-like fungi on Eutypa and Eutypella and with acremonium-to verticillium-like asexual stages. Since then, six additional taxa have joined the group [7][8][9]. ...
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Species of Nectriaceae commonly occur on living and decaying woody substrates, soil, fruitbodies of other fungi, and insects. Some of them are reported as endophytes, opportunistic pathogens of crops and humans, or producers of mycotoxins. To explore the species diversity of the family, specimens from different regions of China were collected and examined. Four novel taxa of Penicillifer, Pseudocosmospora, and Thelonectria were introduced on the basis of morphological characteristics and DNA sequence analyses of combined datasets of the act, ITS, LSU, rpb1, rpb2, tef1, and tub2 regions. Differences between the new species and their close relatives were compared and discussed.
... The genus Pseudocosmospora C.S. Herrera & P. Chaverri, typified by P. eutypellae C.S. Herrera & P. Chaverri, is characterized by scattered to gregarious subglobose to obpyriform, bluntly papillate perithecia that are collapsing laterally when dry, scarlet, KOH+, and usually less than 250 μm in height; cylindrical to narrowly clavate asci containing eight ellipsoidal, yellow-brown, 1-septate and verrucose ascospores; and producing acremonium-to verticillium-like asexual states, cylindrical phialides, and ellipsoidal, ovoid or reniform, smooth, non-septate conidia (Herrera et al. 2013). Some members produce bioactive secondary metabolites which play important roles in biomedicine and agriculture (Lee et al. 2011;Shiono et al. 2016;Nakamura et al. 2019). ...
... & C. Tul. and Eutypella (Nitschke) Sacc., and rarely on Biscogniauxia Kuntze and Hypoxylon Bull. (Herrera et al. 2013;Zeng and Zhuang 2017;Lechat and Fournier 2020). ...
... The isolate CGMCC 3.20177 was shown as a separate lineage and grouped with P. rogersonii C.S. Herrera & P. Chaverri, P. eutypellae, and three unnamed strains. The isolate CGMCC 3.20178 clustered with the unidentified strain MAFF 241531 (Herrera et al. 2013) (MPBP/MLBP/BIPP = 100%/100%/100%) and further grouped with P. rogersonii, which formed a supported terminal branch in Pseudocosmospora (MPBP/MLBP/BIPP = 99%/94%/100%). ...
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... This fungus was assigned to the Nectriaceae based on ascomata changing colour in 3% KOH and lactic acid. The new species described herein is morphologically similar to the known cosmospora-like species occurring on dead or effete stromata of Diatrypaceae and Hypoxylon as recently revised and delimited by GräFEnHAn et al. (2011), HErrErA et al. (2013, LOmbArd et al. (2015), and LEcHAT et al. (2019). Here we explain our morphological, cultural and phylogenetic results leading to the placement of this species in the genus Pseudocosmospora c.S. Herrera & P. chaverri and discuss the features supporting the segregation of P. hypoxylicola from its relatives. ...
... for species with acremonium-like asexual morphs. based on a good correlation between phylogenetic results and the colours of the colonies in culture on PdA, Pseudocosmospora typified by P. eutypellae c.S. Herrera & P. chaverri was introduced for ten cosmospora-like species occurring mostly on stromata of Diatrypaceae, having acremonium-like asexual morphs and producing rosy-buff, pale-luteous to orange or salmon-pink colonies (HErrErA et al., 2013). In contrast, Cosmospora was restricted to species growing on polypores or Xylariaceae, forming green colonies in culture with Cosmospora arxii (Gams) Gräfenhan & Schroers being an exception in having a salmon to saffron colony (HErrErA et al., 2015). ...
... Whenever a reliable identification of the host can be made, it more or less strongly suggests host-specificity of cosmospora-like fungi for a fungal species or genus (HErrErA et al., 2013;. cosmospora-like fungi are rarely reported from Hypoxylon. ...
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The Hypocreales with over one thousand described species have been the repository for all light- to bright-colored, soft-textured, perithecial ascomycetes with a Nectria-type centrum. Rogerson (1970) published a key to the genera in the Hypocreales and accepted over 115 genera with 26 generic synonyms in the order. Since then, 58 genera have been added. For this study all available type specimens of the type species of genera classified in the Hypocreales were examined. Fifty six genera, including six newly described genera with 43 generic synonyms, are accepted in three families, Bionectriaceae fam, nov., Hypocreaceae and Nectriaceae, of the order. Although now considered either part of or closely related to the Hypocreales, neither the Niessliaceae nor the Clavicipitaceae are treated comprehensively in this study. Fourteen genera with two generic synonyms are included in the Niessliaceae and six genera with one generic synonym are placed in the Clavicipitaceae. The remaining 84 genera are excluded from the Hypocreales and redisposed in their appropriate family and order. Genera excluded from the Bionectriaceae, Hypocreaceae, and Nectriaceae are described and illustrated based on their type species. For 16 genera previously placed in the Hypocreales the type specimen was either not located or not sufficient to make a modern taxonomic evaluation of the type species. For each genus the type species and species not recently treated are fully described and documented. A key to species is presented unless a recent key to species in that genus is available. In the Bionectriaceae a new genus, Ochronectria, is introduced for Nectria calami. Nectriella minuta, N. rubricapitula, N. utahensis, Pronectria echinulata, P. pertusariicola, and Protocreopsis viridis are described as new species. 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Crouan, and N. halonata for Charonectria umbelliferarum, are proposed. In the Nectriaceae five new genera are introduced: Albonectria for species related with Nectria rigidiuscula, Haematonectria for the Nectria haematococca complex, Lanatonectria for the Nectria flavolanata-group, Rubrinectria for a species previously known as Nectria olivacea, and Viridispora for teleomorphs of Penicillifer. Cosmospora dingleyae and C. obscura are described as new species. The following new specific combinations are proposed: Albonectria rigidiuscula, A. albosuccinea, and A. verrucosa; Corallomycetella repens and C. jatrophae; Cosmospora aurantiicola, C. biasolettiana, C. camelliae, C. chaetopsinae, C. chaetopsinae-catenulatae, C. chaetopsinae-penicillatae, C. chaetopsinae-polyblastiae, C. chlorina, C. consors, C. digitalicola, C. diminuta, C. diploa, C. episphaeria, C. flammea, C. flavoviridis, C. ganymede, C. geastroides, C. glabra, C. joca, C. jucundula, C. kurdica, C. lasiodiplodiae, C. leptosphaeriae, C. macrochaetopsinae, C. magnusiana, C. meliopsicola, C. metepisphaeria, C. nothepisphaeria, C. papilionacearum, C. peponum, C. pseudepisphaeria, C. pseudoflavoviridis, C. purtonii, C. rickii, C. rishbethii, C. rubrisetosa, C. sansevieriae, C. stilbellae, C. stilbosporae, C. thujana, C. triqua, C. tungurahuana, C. vilior, C. viliuscula, C. wegeliana, and C. xanthostroma; Haematonectria haematococca, H. illudens, H. ipomoeae, H. monilifera, and H. termitum; Lanatonectria flocculenta with anamorph Actinostilbe macalpinei, L. flavolanata, L. mammiformis with anamorph Actinostilbe mammiformis, and L. raripila; Neonectria coccinea and N. galligena; Rubrinectria olivacea; Viridispora penicilliferi, V. alata, V. diparietispora, and V. fragariae; Xenonectriella leptaleae, X. ornamentata, and X. streimannii. In the checklist, some genera are excluded from the families treated here and placed among 19 families in 12 orders of ascomycetes and one basidiomycetous genus. Two genera are uniloculate, discomycetous loculoascomycetes; some have true apothecia and belong in the Helotiales and Pezizales, or are lichenized Lecanorales. Many of these taxa are placed in the Diaporthales and Xylariales (Hyponectriaceae and Thyridiaceae). Genera having immersed ascomata are often difficult to place; they include Charonectria and Hyponectria, now placed in the Hyponectriaceae, Xylariales; and Cryptoleptosphaeria, Cryptonectriella and Schizoparme, now placed in the Diaporthales. Several genera are placed in the Niessliaceae and Clavicipitaceae of the Hypocreales. In this section a new species, Charonectria amabilis, is described, and the new combinations Thyridium ohiense, Charonectria sceptri, Cryptoleptosphaeria gracilis, Cryptonectriella geoglossi, and Thelocarpon citrum, are proposed.