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Coprophilous species of Podospora reported from Brazil are discussed. Thirteen species are recorded for the first time in Northeastern Brazil (Pernambuco) on herbivore dung. Podospora appendiculata, P. australis, P. decipiens, P. globosa and P. pleiospora are reported for the first time in Brazil, while P. ostlingospora and P. prethopodalis are reported for the first time from South America. Descriptions, figures and a comparative table are provided, along with an identification key to all known species of the genus in Brazil.
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Submitted 6 February 2015, Accepted 20 March 2015, Published online 3 April 2015
Corresponding Author:Roger Fagner Ribeiro Melo e-mail rogerfrmelo@gmail.com
Andrew Nicholas Miller e-mail amiller7@illinois.edu
Leonor Costa Maia e-mail leonorcmaia@gmail.com 201
The genus Podospora (Lasiosphaeriaceae, Sordariales) in Brazil
Melo RFR1, Miller AN2 and Maia LC1
1Universidade Federal de Pernambuco, Departamento de Micologia, Centro de Ciências Biológicas, Avenida da
Engenharia, s/n, 50740600, Recife, Pernambuco, Brazil. rogerfrmelo@gmail.com
2 Illinois Natural History Survey, University of Illinois, 1816 S. Oak St., Champaign, IL 61820
Melo RFR, Miller AN, MAIA LC 2015 The genus Podospora (Lasiosphaeriaceae, Sordariales) in
Brazil. Mycosphere 6(2), 201215, Doi 10.5943/mycosphere/6/2/10
Abstract
Coprophilous species of Podospora reported from Brazil are discussed. Thirteen species are
recorded for the first time in Northeastern Brazil (Pernambuco) on herbivore dung. Podospora
appendiculata, P. australis, P. decipiens, P. globosa and P. pleiospora are reported for the first
time in Brazil, while P. ostlingospora and P. prethopodalis are reported for the first time from
South America. Descriptions, figures and a comparative table are provided, along with an
identification key to all known species of the genus in Brazil.
Key words Ascomycota coprophilous fungi taxonomy
Introduction
Podospora Ces. is one of the most common coprophilous ascomycetes genera worldwide,
rarely absent in any survey of fungi on herbivore dung (Doveri, 2008). It is characterized by dark
coloured, non-stromatic perithecia, with coriaceous or pseudobombardioid peridium, vestiture
varying from glabrous to tomentose, unitunicate, non-amyloid, 4- to multispored asci usually
lacking an apical ring and transversely uniseptate two-celled ascospores, delimitating a head cell
and a hyaline pedicel, frequently equipped with distinctly shaped gelatinous caudae (Lundqvist,
1972). Most species are coprophilous, living in dung of many herbivores.
For a historical summary regarding the nomenclature of Podospora, refer to Lundqvist (1972)
and Cain (1934). According to the former author, the genus circumscription is based on: presence
of a basal pedicel usually collapsing, clavate shape of the young ascospores and lack of a thickened
apical ring in the asci. Huhndorf et al. (2004) circumscribed the Lasiosphaeriaceae as a paraphyletic
"lasiosphaeriaceous complex", which included Podospora and 23 additional genera based on
morphology or molecular analyses using partial sequences of the nuclear large-subunit (LSU)
rDNA. Species with swollen agglutinated perithecial hairs and long persistent pedicels, previously
placed in Podospora, are now treated in Schizothecium Corda, which has been supported by
phylogenetic analyses (Huhndorf et al., 2004; Miller & Huhndorf, 2004; Cai et al., 2005; Kruys et
al., 2015). Thus, this genus is not covered in this work.
In Brazil, Podospora, as well as many other Sordariales, has received little attention. Some
collectors have sent material to specialists around the world, such as Roy Franklyn Cain and Nils
Lundqvist, which resulted in few records throughout the literature (Cain, 1962; Lundqvist, 1972).
During a visit in 1998, Richardson (2001) recorded four species: P. argentinensis, P. communis, P.
Mycosphere 6 (2): 201215(2015) ISSN 2077 7019
www.mycosphere.org Article Mycosphere
Copyright © 2015 Online Edition
Doi 10.5943/mycosphere/6/2/10
202
inflatula, and P. pauciseta, along with an unindentified large-spored species from a small number
of dung samples collected at Bonito and Pantanal do Rio Negro, Mato Grosso do Sul.
In the present work, Podospora species recorded in herbivore dung in Brazil from a study of
herbaria and literature are discussed, along with significant recent additions from fresh material
collected in Pernambuco. An identification key to all species recorded in the country is also
provided.
Materials & Methods
In order to access the extent of the diversity of these fungi, dung samples were collected on
farms in Recife, Caruaru and Serra Talhada, and in a Zoological park in Recife, State of
Pernambuco, Northeastern Brazil. Samples were collected in clean plastic bags, taken to the
laboratory and incubated in moist chambers at room temperature (28°C ± 2°C) for at least 75 days
under alternating natural light and dark periods. Material was observed directly from substrata with
the aid of a stereomicroscope, and perithecia were mounted in tap water, Indian ink, Congo red,
lactophenol with cotton blue or Polyvinyl-Lacto-Glycerol for identification under a light
microscope. Species were identified based on macroscopic and microscopic structures according to
Cain (1962), Mirza & Cain (1969), Lundqvist (1972), Bell (1983, 1995), Richardson & Watling
(1997) and Doveri (2004), except in cases where more recent work was available. A careful
literature revision and a survey of national herbaria were performed to access information regarding
older records of Podospora in Brazil. High resolution images of morphological characters were
taken with an Olympus QColor 3 digital camera mounted on an Olympus BX51 microscope
equipped with bright-field and Nomarski interference optics. Permanent slides were mounted and
deposited in URM (Pe. Camille Torrend Herbarium, Universidade Federal de Pernambuco, Recife,
Brazil). For a full list of species synonyms, see Index Fungorum online database (http://
www.indexfungorum.org/). Additional information regarding all records and deposited specimens
along with high quality micrographs are available at the "INCT - Herbário Virtual da Flora e dos
Fungos" database website (http://inct.florabrasil.net/).
Results
Fourteen species were identified from fresh material. An identification key to all known
species of Podospora from Brazil is provided. A comparative table is presented, showing key
morphological characters used to distinguish these species (Table 1). The unverified material was
either unavailable or was too poor to allow an appropriate determination.
Table 1 Characteristics of Podospora species in Brazil
Species
Head cellsize
Asci
Apical cauda
Basal cauda
P. appendiculata
27.5-30 × 12.5-15 µm
8-spored
Single, lash-like, subapical
Single, lash-like, distal
P. australis
57.5 × 25-30.5 µm
4-spored
Single, pleated, subapical
Single, pleated, covering the small
pedicel
P. communis
32.5-40 × 16.5-20 µm
8-spored
Four independent, cylindrical,
subapical
Four independent,
cylindrical,proximal
P. curvicolla
15-17.5 × 8.5-10 µm
Multispore
d
Single, lash-like, fugacious,
apical
Single, lash-like, fugacious, distal
P. decipiens
35-42 × 17.5-22.5 µm
8-spored
Single, cylindrical to doliform,
lamellate, apical
Several, in a lyre-shaped tuft,
proximal
P. fimiseda
52.5-55 × 27.5-30 µm
8-spored
Single, lash-like, with channels,
subapical
Single, lash-like, with channels,
distal
P. globosa
35-40 × 20-25 µm
8-spored
Gelatinous sheath surrounding
the whole spore
Gelatinous sheath surrounding the
whole spore
P. inflatula
27.5-30 × 12.5-15 µm
8-spored
Single, lash-like, subapical
Single, lash-like, distal
P. longicaudata
40-47.5 × 22.5-25 µm
8-spored
Several in one tuft, covering the
head cell
Several in one tuft,distal
P. ostlingospora
52-57.5 × 25-27.5 µm
8-spored
Several in two tufts not
covering the head cell
Single, lash-like, distal
P. pauciseta
35-37.5 × 17.5-19.5
µm
4-spored
Single, cylindrical,subapical
(1) terminal, single, distal; (2)
lateral, 2-4, proximal
P. pleiospora
30-35 × 17.5-22.5 µm
16-32-
spored
Single, lash-like, lamellate,
apical
2-3, lash-like, proximal
203
P. prethopodalis
32-42.5 × 15-20 µm
8-spored
Single, cylindrical, apical
Single, cylindrical, distal
Key to known species of Podospora from Brazil
1. Asci 4-spored .................................................................................................................................. 2
1. Asci with more than 4 spores.......................................................................................................... 3
2. Head cells 35-37.5 × 17.5-19.5 µm. Asci occasionally slightly constricted in the middle. Pedicel
well developed ................................................................................................................. P. pauciseta
2. Head cells 50-57.5 × 25-30.5 µm. Asci not constricted. Pedicel poorly developed ....... P. australis
3. Perithecia with black tubercles near the base of the neck. Apical caudae with longitudinal
lamellae and basal caudae attached to the pedicel base ...................................................................... 4
3. Perithecia without black tubercles near the base of the neck. Above combined characters not
present ................................................................................................................................................. 6
4. Asci 16-spored. Head cells 30-35 × 17.5-22.5 µm ...................................................... P. pleiospora
4. Asci 8-spored. Head cells otherwise .............................................................................................. 5
5. Head cells 35-42 × 17.5-22.5 µm .................................................................................. P. decipiens
5. Head cells 26-34 × 12-20 µm ................................................................................ P. argentinensis
6. Ascospores surrounded by a thin gelatinous sheath. Caudae absent .............................. P. globosa
6. Gelatinous sheath surrounding the spores not observed. Caudae present ..................................... 7
7. Ascospores with more than one apical or basal cauda .................................................................. 8
7. Ascospores with only one apical and one basal cauda ................................................................ 12
8. Asci 64-spored. Ascospores with small caudae attached to the pedicel base ........... P. brasiliensis
8. Asci 8-spored. Caudae not as above .............................................................................................. 9
9. Gelatinous equipment of ascospores composed of four subapical caudae attached to the head cell
and four basal caudae attached to the pedicel distal end .................................................. P. communis
9. Gelatinous equipment of ascospores composed of multiple caudae (more than four), both on the
head cell and on the pedicel ............................................................................................................... 10
10. Cauda split into two tufts, not surrounding the head cell ..................................... P. ostlingospora
10. Cauda tuft single, surrounding the head cell ............................................................................... 11
11. Cauda long, up to 250 µm long ............................................................................ P. longicaudata
11. Cauda very short, up to 3 µm long .............................................................................. P. immersa
12. Perithecia with a few long flexuous, septate hairs just below the neck. Ascospores with long
clavate pedicels, distally swollen ............................................................................. P. dolichopodalis
12. Perithecia with different vestiture. Ascospores with cylindrical pedicels, not distally swollen . 13
13. Perithecia with tufts of setose, agglutinated, non-inflated hairs, concentrated on the neck, evenly
pigmented ........................................................................................................................................ 14
13. Perithecia with non-setose, non-agglutinated hairs, which can be apically inflated or not,
scattered over the entire ascomata, becoming hyaline towards the tip ............................................ 15
14. Asci (128?-)256(-512?)-spored. Pedicels obclavate, non-constricted ....................... P. curvicolla
204
14. Asci 8-spored. Pedicels cylindrical, with a conspicuous constriction, delimiting an inflated basal
part and a slightly inflated apical part ........................................................................ P. prethopodalis
15. Perithecial wall pseudobombardioid ........................................................................................... 16
15. Perithecial wall membranaceous ............................................................................................... 17
16. Head cells 27.5-30 × 12.5-15 µm ....................................................................... P. appendiculata
16. Head cells 52.5-55 × 27.5-30 µm ................................................................................. P. fimiseda
17. Hairs apically inflated .................................................................................................. P. inflatula
17. Hairs not inflated ..................................................................................................... P. longispora
Taxonomy
Podospora appendiculata (Auersw. ex Niessl) Niessl, Hedwigia 22: 156 (1883) Fig. 1, AC
Basionym: Sordaria appendiculata Auersw. ex Niessl (1870)
Perithecia scattered to isolated, superficial, ovoid to subellipsoid, with unclear delimitation
between body and neck, light brown to dark brown, darkening towards the neck, 690-775 × 400-
550 µm, covered by setose, rigid, sparse, septate, cylindrical hairs, with obtuse ends, olivaceous
brown, becoming lighter towards the distal tip, finally hyaline at the apex, 2.5-3.5 µm thick, slightly
thicker at the distal tip, up to 5 µm, 62.5-75 µm long. Neck carbonaceous, dark brown to black,
smooth, glabrous. Peridium pseudobambardioid, translucid, 4-layered, outermost layer textura
angularis, with flattened cells, thin-walled, 4.5-15 µm. Paraphyses intertwined with the asci,
ventricose, persistent or collapsing in maturity. Asci 8-spored, clavate, broader at the center, 200-
230 × 32.5-37.5 m, usually persistent, rounded at apex, with a long stipe. Ascospores irregularly
biseriate, hyaline and fusoid to clavate at first, smooth, becoming transversely septate at maturity;
head cell ellipsoid, slightly umbonate at apex, truncate at base, equilateral, aseptate, dark brown to
black, with an apical germ pore, 27.5-30 × 12.5-15 µm, smooth; pedicel cylindrical to obconical,
usually collapsing, 12.5-17.5 × 5-6.5 µm; apical cauda single, lash-like, attached to the apical or
subapical part of the head cell, usually covering the germ pore, 45-47.5 × 5.5-7.5 µm, smooth,
persistent or collapsing in mounting; basal cauda single, morphologically similar to the apical
caudae, slightly thinner, attached to the base of the pedicel, 45-47.5 ×3.5-5 µm, usually collapsing
in mounting.
Habitat Recorded on dung of many herbivores.
Known distribution Central America (Costa Rica), Europe (Belgium, Czech Republic,
Denmark, England, Finland, France, Germany, Iceland, Ireland, Italy, Norway, Scotland, Sweden),
North America (Canada, Greenland, USA) and Oceania (Australia, New Zealand). This is the first
record from Brazil (see the discussion under "Podospora longispora").
Material examined Brazil, Pernambuco, Recife, Universidade Federal Rural de Pernambuco
(UFRPE), on horse dung, 31 Jul 2012, R.F.R. Melo (URM86729a, 86729b, 86729c), 29 Nov 2011,
R.F.R. Melo (URM86730a, 86730b), on cattle dung, 28 Jan 2013, R.F.R. Melo (URM86731).
Notes Podospora appendiculata was commonly found in samples collected in Recife, but
absent in collections from the semi-arid region. It is distinguished by its superficial perithecia with
a pseudobombardioid wall and stiff, septate perithecial hairs with a hyaline apex. Perithecia of P.
fimiseda show a similar morphology regarding the wall and hairs, but can be easily differentiated
by the larger ascospore head cells (48-60 × 27-31 µm). Podospora inflatula and P. australis are
also similar, but perithecia are clearly separated into a body and neck and have membranaceous
walls. Podospora australis has perithecial hairs solely on the neck and four ascospores per ascus,
with head cells considerably larger.
Podospora australis (Speg.) Niessl, Hedwigia 22: 156 (1883) Fig. 1, C
Basionym: Hypocopra australis Speg., Anal. Soc. cient. argent. 10(3): 137 (1880)
205
Perithecia scattered to isolated, immersed to semi-immersed, obpyriform, light brown to
olivaceous brown, darkening towards the neck to finally black, 650-850 × 390-440 µm, sparsely
covered by flexuous, septate, smooth hairs, light brown in color, 2-2.5 µm thick. Neck cylindrical,
carbonaceous, dark brown to black, smooth, opaque, 190-200 × 180-190 µm, adorned with smooth,
simple or branched, rigid, cylindrical, septate, hairs, acute to obtuse tipped, light brown, becoming
lighter towards the distal end to finally hyaline, 2.5-3.5 µm thick, extending for up to 60 µm.
Peridium membranaceous, translucid, 3-layered, outermost layer textura angularis, with flattened
cells, thin-walled, 5-12.5 µm long. Paraphyses interspersed with the asci, filiform, usually
persistent. Asci 4-spored, cylindrical-clavate, 260-300 × 37.5-45 µm, usually persistent, with a
rounded apex and a long stipe. Ascospores uniseriate, at first hyaline, clavate to saccate, aseptate,
smooth, later becoming swollen in the upper portion; head cell ellipsoid, with apical germ pore and
rounded base, equilateral, aseptate, luteous, light brown, dark brown or black, 57.5 × 25-30.5 µm,
smooth; pedicel absent or, when present, reduced, diminished, obconical, up to 2.5 µm in length,
hyaline or slightly pigmented; apical cauda cylindrical, gradually decreasing in diameter towards
the apex, provided with multiple internal channels, marked by segments along its length, pleated,
adhered to the subapical part of the head cell, not covering the germ pore, 175-215 × 7.5-10 µm,
persistent; basal cauda symmetrical, covering the pedicel when present, devoid of internal channels,
pleated, 150-210.5 ×10-12.5 µm, persistent.
Habitat Recorded on dung of many herbivores.
Known distribution Despite being worldwide, this is the first record for Brazil.
Material examined Brazil, Pernambuco, Caruaru, Instituto Agronômico de Pernambuco
(IPA), on goat dung, 30 Sep 2011, R.F.R. Melo (URM86732a, 86732b).
Notes Podospora australis is unique in having 4-spored asci bearing ascospores with large
head cells (50-57.5 ×25-30.5 µm) with a rounded base, unlike the truncated base usually observed
in other Podospora species, reduced (or absent) pedicels, along with persistent and pleated caudae.
It is distinguished from other 4-spored species such as: Schizothecium nanum and S. tetraspora by
the size of the ascospores and by the absence of inflated swollen hairs on the perithecium; P.
inequalis by the presence of a gelatinous apparatus in the ascospores; and P. pauciseta by the
reduced state of the pedicel. Doveri (2004) presents an identification key for the 4-spored members
of Podospora and Schizothecium.
Podospora communis (Speg.) Niessl, Hedwigia 22: 156 (1883) Fig. 1, DF
Basionym: Hypocopra communis Speg., Anal. Soc. cient. argent. 10(1): 14 (1880)
Perithecia scattered to isolated, semi-immersed, obpyriform, 770-800 × 370-450 µm; body
dark brown to black, glabrous or covered by slender and flexuous hairs, olivaceous brown, 2-2.5
µm thick. Neck cylindrical, carbonaceous, blackening near the apex, opaque, 250-360 × 140-150
µm, smooth, glabrous, usually markedly extending towards the light, becoming bend to crooked.
Peridium membranaceous, subopaque, 3-layered, outermost layer textura angularis, with thick-
walled cells, 5-9.5 µm long. Paraphyses interspersed with the asci, ventricose, persistent or
collapsing. Asci8-spored, clavate, 240-255 × 45-55 µm, usually persistent, with narrowed,
truncated apex and a long stipe. Ascospores biseriate, at first hyaline, clavate, aseptate, smooth,
later becoming swollen in the upper portion; head cell ellipsoid, slightly umbonate at apex, truncate
at base, aseptate, reddish brown to dark brown, with an apical germ pore, 32.5- 40 × 16.5-20 µm,
smooth; pedicel cylindrical, persistent or collapsing, 6.5-7.5 × 25-28.5 µm; apical gelatinous
equipment composed of four hyaline, cylindrical apical caudae, gradually decreasing in diameter
towards the apex, attached to the upper part of the head cell, 20-25 × 4-5 µm, smooth, persistent or
collapsing; basal gelatinous equipment composed of four hyaline caudae, similar to the apical
caudae in morphology, but shorter, attached to the base of the pedicel, 2-5 × 1-3 µm, persistent or
collapsing.
Habitat Recorded on dung ofmany herbivores.
Known distribution Worldwide.
206
Material examined Brazil, Pernambuco, Caruaru, Instituto Agronômico de Pernambuco
(IPA), on cattle dung, 07 Oct 2011, R.F.R. Melo (URM86733), 03 Mar 2012, R.F.R. Melo
(URM86734), 03 Sep 2012, R.F.R. Melo (URM86735), Serra Talhada, Instituto Agronômico de
Pernambuco (IPA), 30 Apr 2013, R.F.R. Melo (URM86736), Recife, Universidade Federal Rural
de Pernambuco (UFRPE), on cattle dung, 02 May 2012, R.F.R. Melo (URM86737).
Notes As its epithet implies, Podospora communis was found to be common and
widespread in herbivore dung in Pernambuco, mainly occurring on the dung of domesticated
animals. Itis recognized byits glabrous perithecium, strongly phototropic neck and ascospores with
a gelatinous apparatus composed of four subapical caudae at the head cell and four caudae at the
end of the pedicel. It differs from other multicaudate species such as P. immersa and P.
ostlingospora by having only four apical and four basal caudae.
Podospora curvicolla (G. Winter) Niessl, Hedwigia 22: 156 (1883) Fig. 1, HJ
Basionym: Sordaria curvicolla G. Winter 1871
Perithecia scattered to isolated, semi-immersed to immersed, obpyriform, light brown,
becoming darker and finally black at the neck, 400-560 × 380-450 µm, covered by long, flexuous,
brown hairs, 2-3 µm thick. Neck cylindrical, 100-170 × 80-150 µm, with tufts of long, cylindrical,
setose, agglutinated hairs, extending up to 400 µm long. Peridium membranaceous, opaque to
subopaque, without clear distinction between layers, outermost layer textura angularis. Asci (128-
?)256(?-512)-spored, saccate to clavate, stipitate, 200-320 × 70-115 µm. Ascospores multiseriate,
hyaline, clavate or rod-shaped when immature, later becoming swollen in the upper part; head cell
ellipsoid, truncated at base, with an apical germ pore, olivaceous brown to dark brown, 15-17.5 ×
8.5-10(-11.5) µm, equilateral; pedicel obclavate, 5-7 × 2-3 µm, usually collapsing; gelatinous
equipment comprised of single lash-like caudae on the apical and basal ends of the spore,
fugacious, hardly observable in the material examined.
Habitat Recorded on dung of many herbivores.
Known distribution Europe (Belgium, Germany, Hungary, Italy, Poland, Scotland,
Sweden), North America (Canada, USA), South America (Brazil, see ‘Notes’).
Material examined Brazil, Pernambuco, Recife, Universidade Federal Rural de Pernambuco
(UFRPE), on horse dung, 13 Mar 2012, R.F.R. Melo (URM86738, 86739).
Notes Among the multispored species of Podospora, P. curvicolla differs from members of
Schizothecium in having setose hairs around the neck, P. araneosa by having ascospores with larger
head cells (8.5-10 × 15-17.5 µm), and P. setosa and P. bifida by having perithecial hairs forming
tufts. Lundqvist (1972) mentions that Batista & Pontual had described and identified Philocopra
coprophila from the material that, although he was not able to examine, he believed to be a
specimen of P. curvicolla and recommends its position as nomem dubium. Podospora curvicolla
was recorded in Pernambuco, Brazil, by Melo et al. (2012).
Podospora decipiens (G. Winter ex Fuckel) Niessl, Hedwigia 22: 156 (1883) Fig. 1, KM
Basionym: Sordaria decipiens G. Winter, Abh. naturforsch. Ges. Halle 13(1): 28 (1873)
Perithecia scattered, semi-immersed to superficial, obpyriform, olive brown, becoming darker
and finally black at the neck, 500-950 × 300-490 µm, glabrous or covered by slender, flexuous
hairs, olive brown, 1.5-2.5 µm thick. Neck cylindrical, carbonaceous, opaque, 200-320 × 150-160
µm, smooth, adorned throughout its length by dark brown to black thick-walled tubercles,
numerous, obtuse, straight or sinuous, 10-20 × 4.5-5 µm, densely clustered in the neck, giving it a
strongly carbonaceous texture, becoming gradually and progressively sparse in the upper body,
absent in the lower part of the ascoma. Peridium membranaceous, subopaque, 3-layered, outermost
layer textura angularis, with thick-walled cells, 7.5-12.5 µm long. Paraphyses interspersed with the
asci, ventricose, usually persistent. Asci 8-spored, clavate, 300-330 × 45-47.5 µm, with narrowed
truncate apex and a long stipe. Ascospores biseriate, hyaline when immature, later becoming
swollen in the upper portion; head cell ellipsoid, slightly apiculated at apex, truncate at base,
aseptate, reddish brown to dark brown, with a subapical germ pore, 35-42 × 17.5-22.5 µm, smooth;
207
pedicel cylindrical, persistent or collapsing, 50-60 × 5-7.5 µm, with a slight distal swelling in some
ascospores; apical caudae cylindrical to doliform, lamellate, attached to the apex of the head cell,
20-25 × 10-12.5 µm, smooth, persistent or collapsing, not covering the germ pore; basal caudae
numerous, in a lyre-shaped tuft, attached to the proximal end of the pedicel, up to 10-12.5 µm in
length, usually collapsing.
Habitat Recorded on dung of many herbivores.
Known distribution Despite being worldwide, this is the first record for Brazil.
Material examined Brazil, Pernambuco, Recife, Universidade Federal Rural de Pernambuco
(UFRPE), on goat dung, 05 May 2012, R.F.R. Melo (URM86740a, 86740b).
Notes Podospora decipiens, a commonly recorded species in herbivore dung, is similar to
P. pleiospora and P. myriaspora, especially due to the presence of black tubercles on the neck and
the similarity of the ascospore caudae. It differs from the former by the number of ascospores per
ascus (eight) and from the latter by the significantly larger head cells (17.5-22.5 × 35-40 µm in P.
myriaspora). Lundqvist (1972) did not observe any material of P. decipiens from the tropics.
Podospora fimiseda (Ces. & De Not.) Niessl, Hedwigia 22: 156 (1883) Fig. 1, NO
Basionym: Sordaria fimiseda Ces. & De Not., Comm. Soc. crittog. Ital. 1: 226 (1863)
Perithecia gregarious to isolate, superficial, obpyriform, 690-930 × 510-580 µm, olive brown,
becoming darker and finally black at the neck, sparsely covered by stiff, septate, cylindrical hairs,
hairs obtuse ended, thick-walled, olivaceous brown, becoming lighter toward the distal end to
finally hyaline at the tip, 3-3.5 µm thick, up to 85 µm long. Neck cylindrical, carbonaceous,150-205
× 200-225.5 µm. Peridium pseudobombardioid, subopaque, coriaceous, 4-layered, outermost layer
textura angularis,cells flattened, thin-walled. Paraphyses interspersed with the asci, ventricose,
persistent or collapsing. Asci 8-spored, clavate, wider at the medial portion of the sporiferous part,
385-405 × 50-60 µm, usually persistent, with truncated apex and a long stipe. Ascospores
irregularly biseriate, hyaline when immature, fusoid to clavate, later becoming swollen in the upper
portion; head cell ellipsoid, slightly umbonateat apex, truncate at base, equilateral, aseptate, reddish
brown to dark brown, with an apical germ pore, 52.5-55 × (25-)27.5-30 µm, smooth; pedicel
cylindrical, persistent, 30-35 × 5-7.5 µm, slightly thickened in the distal end, 7.5-8.5 µm; apical
caudae lash-like, with internal channels; basal caudae similar in morphology, shorter and slightly
thinner, attached to the base of the pedicel.
Habitat Recorded mainly on dung of domestic herbivores.
Known distribution Africa (Liberia), Asia (China), Central America (Puerto Rico), Europe
(Austria, Belgium, Denmark, Finland, France, Germany, Hungary, Italy, Norway, Poland,
Romania, Russia, Scotland, Spain), South America (Brazil, Chile, Venezuela), North America
(USA, Canada), Oceania (Australia, New Zealand).
Material examined Brazil, Pernambuco, Caruaru, Instituto Agronômico de Pernambuco
(IPA), on horse dung, 16 Oct 2011, R.F.R. Melo (URM86741).
Notes Podospora fimiseda may initially be confused with P. appendiculata due to the
pseudobombardioid peridium and perithecia covered with hyaline-tipped hairs. However, the
former differs markedly from the latter by having perithecia with a clear delineation between the
body and neck and ascospores with head cells significantly larger (52.5- 55 × 27.5-30 µm).
Podospora australis possesses hairs with similar morphology, but has four ascospores per ascus,
reduced or absent pedicel and different caudae.
Podospora globosa (Massee & E.S. Salmon) Cain, Can. J. Bot. 40: 460 (1962) Fig. 1, P
Basionym: Sordaria globosa Massee & E.S. Salmon, Ann. Bot., Lond. 15: 334 (1901)
Perithecia scattered, partially immersed on dung, obpyriform, light brown to olivaceous,
becoming darker and finally black at the neck, 600-700 500-600 m. Neck roughly cylindrical,
opaque, glabrous. Peridium membranaceous, becoming carbonaceous, subopaque, 3-layered,
outermost layer textura angularis, cells 7.5-12.5 µm. Asci 8-spored, clavate, rounded at apex, with
a short stipe, 270-350 30-40 m. Ascospores biseriate, hyaline when immature, fusoid to clavate,
208
later becoming swollen in the upper portion; head cell ellipsoid, truncate at base, with an apical
germ pore, dark brown to black, (32.5-)35-40 (17.5-)20-25 m, smooth, equilateral; pedicel
cylindrical, (21-)22.5-37.5 5-7 m, usually persistent; gelatinous sheath surrounds the entire
spore, ~2.5-4.5 µm thick, hyaline.
Habitat Recorded on dung of many herbivores.
Known distribution Europe (England, Spain, Sweden) and North America (Canada). This is
the first record for South America.
Material examined Brazil, Pernambuco, Recife, Horto Zoobotânico do Parque Estadual
Dois Irmãos, on llama dung, 17 Mar 2010, R.F.R. Melo (URM82322), on deer dung, 22 Mar 2010,
R.F.R. Melo (URM82323, 82324).
Fig. 1 Podospora appendiculata. (A) Perithecium (bar = 150 µm), (B) rigid, hyaline-tipped
perithecial hairs (bar = 10 µm). Podospora australis. (C) Ascus with four mature ascospores (bar =
25 µm). Podospora communis. (D) Perithecium (bar = 100 µm), (E) mature ascus (bar = 10 µm),
(F) ascospore (bar = 12.5 µm). Podospora curvicolla. (G) Ascospore (bar = 7.5 µm), (H)
perithecium (bar = 100 µm), (I) immature ascus (bar = 35 µm), (J) mature ascus (bar = 35 µm).
209
Podospora decipiens. (K) Perithecium (bar = 100 µm), (L) ascospore (bar = 20 µm), (M) immature
ascus (bar = 25 µm). Podospora fimiseda. (N) Perithecium (bar = 100 µm), (O) ascospore (bar = 20
µm). Podospora globosa. (P) Perithecium (bar = 100µm).
Notes Podospora globosa is an atypical member of the genus by having a conspicuous
gelatinous layer surrounding the entire ascospore, similar to that seen in Sordaria. The long pedicel
(22.5-37.5 µm), glabrous perithecial neck and the biseriate organization of ascospores are other
important distinguishing characters of this species.
Podospora inflatula Cain, Can. J. Bot. 40: 454 (1962) Fig. 2, AB
Perithecia scattered to gregarious, superficial, obpyriform, 550-650 × 390-410 µm, olive
brown, becoming darker to finally black towards the neck, covered along its entire length by sparse,
septate, cylindrical, setose hairs, thick-walled, becoming clearly inflated at the apex, initially
olivaceous brown, becoming lighter towards the distal end to finally hyaline, 3.5-5 µm thick near
the base, swelling up to 8-8.5 mm thick at the apex, extending up to 70 µm long. Neck cylindrical,
carbonaceous, opaque, 90-120 × 100-110 µm, smooth, glabrous. Peridium pseudobombardioid,
subopaque, 3-layered, outermost layer textura angularis, cells flattened, thin-walled. Paraphyses
ventricose, usually persistent. Asci 8-spored, clavate, slightly wider at the medial portion of the
sporiferous region, 150-200 × (20-)25-32.5 µm, usually persistent, narrowed and rounded above,
with a long stipe. Ascospores irregularly biseriate, hyaline when immature, fusoid, later becoming
swollen in the upper portion; head cell ellipsoid, slightly umbonate at apex, truncate at base,
equilateral, asseptate, reddish brown to dark brown, with an apical germ pore, 27.5-30 × 12.5-15
µm, smooth; pedicel cylindrical, persistent or collapsing, 25-30 × 5-6 µm; apical caudae lash-like,
with internal channels, attached to the subapical portion of the head cell, 20-35 × 5-5.5 µm; basal
caudae similar in morphology, shorter and thinner, adhered to the base of the pedicel,~1-1.5 mm
diam. Habitat Recorded on horse dung.
Known distribution Central America (Mexico), Oceania (Society Islands) and South
America (Brazil).
Material examined Brazil, Pernambuco, Recife, Universidade Federal Rural de Pernambuco
(UFRPE), on horse dung, 12 Mar 2012, R.F.R. Melo (URM86742), 13 Mar 2012, R.F.R. Melo
(URM86743a, 86743b).
Notes Podospora inflatula possesses perithecial hairs that are similar to those in P. fimiseda
and P. apendiculata, but can be distinguished by the capitate apices. It differs from P. fimiseda by
having ascospores with shorter head cells (27.5- 30 × 12.5-15 vs. 52.5-55 × 27.5-30 µm), and from
P. appendiculata by having obpyriform perithecia, with a clear distinction between the body and
neck. It was recorded on incubated cattle dung collected in Rio Grande do Sul, Brazil by Cain
(1962).
Podospora longicaudata (Griffiths) Cain, Can. J. Bot. 40: 460 (1962) Fig. 2, CD
Basionym: Pleurage longicaudata Griffiths, Mem. Torrey bot. Club 11: 81 (1901)
Perithecia scattered to gregarious, semi-immersed, obpyriform, 610-650 × 500-560 µm,
olivebrown, becoming black towards the neck, glabrous or covered by slender, flexuous, olivaceous
hairs, 2-3 µm thick. Neck cylindrical, carbonaceous, black near the ostiole, opaque, 150-175 × 130-
145 µm, smooth, glabrous. Peridium membranaceous, subopaque, 3-layered, outermost layer
textura angularis, with thin-walled cells, 4.5-10 µm long. Paraphyses interspersed with asci,
ventricose, usually collapsing. Asci 8-spored, clavate, 250-295 × 30-42.5 µm, with narrowed
truncated apex and a long stipe. Ascospores biseriate, hyaline when immature, clavate, later
becoming swollen in the upper portion; head cell ellipsoid, slightly umbonate at apex, slightly
truncate at base, aseptate, reddish brown to dark brown, with an apical germ pore, 40-47.5 × 22.5-
25 µm, smooth; pedicel clavate to cylindrical, persistent or collapsing, 35-40 × 6.5-7.5 µm; apical
caudae numerous, hyaline, slender, attached to the apex of the head cell, assuming a conical shape,
210
extending up to 250 µm long, smooth, persistent or collapsing; basal caudae similar in morphology,
attached to the base of the pedicel, slender, 2-3 mm thick, persistent or collapsing.
Habitat Recorded on dung of domestic herbivores.
Known distribution Asia (Pakistan), Central America (Mexico, Puerto Rico), North
America (Canada) and South America (Brazil).
Material examined Brazil, Pernambuco, Caruaru, Instituto Agronômico de Pernambuco
(IPA), on goat dung, 07 Oct 2011, R.F.R. Melo (URM86744).
Notes Podospora longicaudata is distinguished by its glabrous perithecia with ascospores
adorned by two conical crowns composed of several caudae at the head cell. Podospora communis,
which also has multiple basal and apical caudae, differs from P. longicaudata in that the caudae do
not form mucilaginous crowns on the ascospores. Mirza & Cain (1969) emphasized the great
variation in the ascospore measurements in this species (40-47.5 × 22.5-25 µm in the examined
material).
Podospora ostlingospora Cain, Can. J. Bot. 40: 456 (1962) Fig. 2, E
Perithecia scattered, immersed, obpyriform, 870-900 × 650-700 µm, faintly reddish brown,
glabrous or covered by slender, flexuous, dark brown hairs, 2-2.5 µm thick. Neck cylindrical,
carbonaceous, black near the ostiole, opaque, 230-250 × 150-155 µm, smooth, glabrous. Peridium
membranaceous, subopaque, outer layer textura angularis, cells thick-walled. Paraphyses
interspersed with asci, ventricose, usually collapsing. Asci 8-spored, clavate, 285-340 × 50-60 µm,
with truncated narrowed apex and a long stipe. Ascospores biseriate, hyaline when immature,
clavate, later becoming swollen in the upper portion; head cell fusoid to long-ellipsoid, slightly
umbonate at apex, truncate at base, aseptate, reddish brown to dark brown, with an apical germ
pore, 52-57.5 × 25-27.5 µm, smooth to slightly rough; pedicel subclavate to cylindrical, hyaline or
rarely pigmented, persistent or collapsing, 27.5-32.5 × 6.5-7.5 µm, usually with slight distal
swelling; apical caudae numerous, split into two tufted tails, usually united, attached to the apex of
the head cell, assuming a roughly conical shape, smooth, persistent or collapsing; basal cauda
single, cylindrical, smooth, attached to the base of the pedicel, usually collapsing.
Habitat Recorded on goat and horse dung.
Known distribution Central America (Mexico). This is the first record from South America.
Material examined Brazil, Pernambuco, Serra Talhada, Instituto Agronômico de
Pernambuco (IPA), on horse dung, 09 Jan 2012, R.F.R. Melo (URM86745), Recife, Universidade
Federal Rural de Pernambuco (UFRPE), on horse dung, 11 Mar 2013, R.F.R. Melo (URM86746).
Notes Podospora ostlingospora differs from other species of the genus by the head cell
shape, fusoid to long-ellipsoid, and size (52-57.5 × 25-27.5 µm), and by the two tufts of caudae at
the apical end. Podospora longicaudata differs in having a single tuft of caudae. Podospora
ostlingospora was first described by Cain (1962) from material growing on incubated burro dung
collected in San Luis Potosi, Mexico, in 1960.
Podospora pauciseta (Ces.) Traverso, Fl. ital. crypt., Fungi 2, 2: 431 (1907) Fig. 2, FH
Basionym: Sphaeria pauciseta Ces., Bot. Ztg. 10: 396 (1852)
Perithecia usually gregarious, partially immersed to superficial, obpyriform, 630-700 × 350-
420 µm, reddish brown to dark brown, glabrous or rarely covered by brown slender, flexuous, hairs,
olivaceous brown to golden, 2-2.5 µm thick. Neck cylindrical, rarely conical, carbonaceous, black
near the ostiole, opaque, 150-185 × 110-140 µm, glabrous or adorned on one side only by a tuft of
non-inflated stiff,hairs, long, setose, erect, septate, with acute tips, olivaceous brown, 2.5-3.5 mm
thick, extending up to 100 µm. Peridium membranaceous, opaque, outer layer textura angularis,
cells thick-walled, 5-12.5 µm long. Paraphyses interspersed with the asci, usually surpassing them
in length, ventricose, persistent or collapsing. Asci 4-spored, clavate, occasionally with a medial
constriction, with narrowed truncated apex and a short stipe, 200-250 × 25-27.5 µm, usually
persistent, occasionally with aborted spores that are significantly smaller than the others.
Ascospores irregularly uniseriate, hyaline when immature, clavate, becoming swollen in the upper
211
portion; head cell ellipsoid, slightly umbonate at apex, truncate at base, dark brown to black,
aseptate, with an apical germ pore, 35-37.5 × 17.5-19.5 µm, smooth; pedicel cylindrical, usually
collapsing, 3-3.5 × 15-17.5 µm; apical cauda cylindrical, gradually decreasing in diameter towards
the apex, 70-75 × 6.5-9 µm, usually collapsing; basal caudae of two kinds: (1) terminal cauda,
similar to thesingle apical cauda, attached to the base of the pedicel, 60-70 × 2.5-5 µm, collapsing;
(2) lateral caudae, 2-4, attached to the proximal end of the pedicel, ~1.5-2 µm thick, extending up to
20 µm long.
Habitat Recorded on dung ofmany herbivores.
Known distribution Worldwide.
Material examined Brazil, Pernambuco, Caruaru, Instituto Agronômico de Pernambuco
(IPA), on horse dung, 30 Nov 2011, R.F.R. Melo (URM86747), Serra Talhada, Instituto
Agronômico de Pernambuco (IPA), on horse dung, 08 Sep 2011, R.F.R. Melo (URM86748),
Recife, Universidade Federal Rural de Pernambuco (UFRPE), on cattle dung, 03 Apr 2012, R.F.R.
Melo (URM86749).
Notes Podospora pauciseta is a common component of the coprophilous mycobiota of
Pernambuco, being the dominant species in this study sampled throughout the state. It can be
distinguished from P. australis, which also has 4-spored asci, bythe smaller head cells (35-37.5 ×
17.5-19.5 µm). Due to the easily collapsing gelatinous appendages, it can be confused with Arnium
arizonense, which also has four ascospores per asci, perithecia of similar size and tufts of erect
hairs, but is distinguished by the smaller head cells.
Podospora pleiospora (G. Winter) Niessl, Hedwigia 22: 156 (1883) Fig. 2, IL
Basionym: Sordaria pleiospora G. Winter, Abh. naturforsch. Ges. Halle 13(1): 13 (1873)
Perithecia usually gregarious, partially immersed to immersed, obpyriform, 750-930 × 430-
550 µm, olive-brown to dark brown, glabrous or covered by slender, flexuous hairs, olivaceous,
1.5-2 µm thick. Neck cylindrical, carbonaceous, black near the ostiole, opaque, 195-250 × 140-185
µm, smooth, glabrous, adorned throughout its length by several dark brown to black thick-walled
tubercles, obtuse, straight or sinuous, simple, 17 5-20 × 3-4.5 µm, densely clustered in the neck
giving it a strongly carbonaceous texture, becoming gradually and progressively sparse in the upper
body, absent in the part of the perithecium. Peridium membranaceous, subopaque, 3-layered,
outermost layer textura angularis, with thick-walled cells, 5-10 µmlong. Paraphyses interspersed
with the asci, ventricose, usually persistent. Asci 16-32-spored, saccate to clavate, 275-300 × 60-
67.5 µm, usually collapsing, with narrowed truncated apex and a long stipe. Ascospores
multiseriate, hyaline when immature, clavate, becoming swollen in the upper portion; head cell
ellipsoid, slightly umbonate at apex,truncate at base, aseptate, reddish brown to dark brown, with an
apical germ pore, 30-35 × 17.5-22.5 µm, smooth; pedicel cylindrical, persistent or collapsing, 5-7.5
× 37.5-45 µm, with distal swelling in some ascospores; apical caudae lash-like, lamellate, head cell
20-25 × 4-5 µm, smooth, persistent or collapsing, not covering the germ pore; basal cauda 2-3,
similar to the apical caudae in morphology, but shorter, attached to the proximal end of the pedicel,
7.5-10 µm long, usually collapsing.
Habitat Recorded on dung of many herbivores.
Known distribution Asia (Taiwan), Central America (Mexico), Europe (Czech Republic,
Germany, Denmark, Greece, Hungary, Italy, Norway, Poland, Romania, Sweden, Switzerland
Spain, United Kingdom) North America (Canada, USA), Oceania (Australia, New Zealand) and
South America (Argentina, Chile).This is the first record from Brazil.
Material examined Brazil, Pernambuco, Recife, Universidade Federal Rural de Pernambuco
(UFRPE), on horse dung, 15 Feb 2012, R.F.R. Melo (URM86750a, 86750b), 23 Mar 2012, R.F.R.
Melo (URM86751).
Notes Three species of Podospora have black tubercles on the perithecia and lamellate
gelatinous caudae. Among them, P. pleiospora can be easily distinguished from P. decipiens, which
has only eight ascospores per ascus, and from P. myriaspora, which has 64 ascospores per ascus.
These species may also be distinguished based on the size of the head cell: P. decipiens (36-42 ×
212
20-22 µm) > P. pleispora (30-35 × 17.5-22.5 µm) > P. myrisaspora (25-34 × 14-19 µm). Some
ascospores of P. pleispora had pedicels with obvious distal swelling.
Podospora prethopodalis Cain, Can. J. Bot. 40: 458 (1962) Fig. 2, M
Perithecia scattered to gregarious, partially immersed to superficial, subglobose to
obpyriform, 320-520 × 225-365 µm, light brown to brown, abundantly covered by slender, septate,
flexuous hairs, olivaceous to golden, 2-2.5 µm thick. Neck cylindrical, carbonaceous, black near the
ostiole, opaque, 100-120 × 90-115 µm, smooth, adorned by tufts of non-inflated, septate, straight,
simple, setose hair, olivaceous brown, ~1.5-2 µm thick, extending up to 300 µm long. Peridium
membranaceous, subopaque, 3-layered, outermost layer textura angularis, with thick-walled cells,
5-12.5 µm long. Paraphyses interspersed with the asci, ventricose, persistent or collapsing. Asci 8-
spored, clavate, 190-215 × 37.5-50 µm, with truncated apex and a long stipe. Ascospores biseriate,
hyaline when immature, clavate, becoming swollen in the upper portion; head cell long-ellipsoid to
fusoid, slightly umbonate at apex, truncate at base, aseptate, reddish brown to dark brown, with a
subapical germ pore, 32-42.5 × 15-20 µm, smooth, usually symmetrical; pedicel cylindrical, usually
collapsing, notably inflated medially and slightly distally, 5-7.5 × 22.5-30 µm; apical cauda single,
cylindrical, attached to the apex of the head cell, 35-42.5(-57.5) × 6-8.4 µm, smooth, persistent or
collapsing; basal cauda single, similar to the apical cauda in morphology, but thicker, attached to
the base of the pedicel, 18.5-35 × 2.5-7.5 µm, usually collapsing.
Habitat Recorded on goat and horse dung.
Known distribution Africa (Kenya), Asia (Pakistan, Taiwan), Central America (Mexico),
North America (Canada), Oceania (Australia, Society Islands). This is the first record from South
America.
Material examined Brazil, Pernambuco, Caruaru, Instituto Agronômico de Pernambuco
(IPA), on goat dung, 15 Jun 2012, R.F.R. Melo (URM86752), Serra Talhada, Instituto Agronômico
de Pernambuco (IPA), on horse dung, 03 May 2012, R.F.R. Melo (URM86753), Recife,
Universidade Federal Rural de Pernambuco (UFRPE), on horse dung, 13 Jun 2012, R.F.R. Melo
(URM86754).
Notes The tufts of agglutinated perithecial hairs and ascospores with medially inflated
pedicels distinguishes Podospora prethopodalis. It was first described by Cain (1962) from material
growing on incubated horse dung collected in Toronto in 1956. The currently known records
suggest a worldwide distribution
Unverified and doubtful species
Podospora argentinensis (Speg.) J.H. Mirza & Cain, Can. J. Bot. 47(12): 2008 (1970) [1969]
Pleurage argentinensis (Speg.) C. Moreau, Encyclop. Mycol. 25: 252 (1954)
Sordaria argentinensis Speg., Anal. Mus. nac. Hist. nat. B. Aires 23: 49 (1912)
This species, whose material was described from Argentina, Mexico and USA, was found in
Brazil by Richardson (2001) and sent to his personal collection (MJR 64/98). The original material
was identified as P. decipiens, which indeed shows P. decipiens-like perithecia but, according to
the author, with smaller ascospores (29-32 × 16 μm). The caudae at the tip of the pedicel were not
observed. Although this material was not examined in this study, based on the author's description,
it may be a good species that should be included in the list of Podospora species recorded in Brazil.
Podospora brasiliensis Cain, Can. J. Bot. 40: 447 (mistyped as 449 in some databases) (1962)
Along with other new species and combinations of Podospora, Roy Franklyn Cain (1962)
proposed P. brasiliensis from material on incubated cattle and sucuri (Eunectes murinus) dung
collected by Alfons Theobald in São Leopoldo, Rio Grande do Sul, Brazil in 1936 (TRTC 35445),
and by Augusto Chaves Batista in Pernambuco in 1957 (TRTC 35593), respectively. These
specimens show some resemblance to P. pleiospora regarding the multispored asci and the small,
black tubercles on the neck of the perithecia, but can be distinguished by the smaller ascospores.
213
Fig. 2 Podospora inflatula. (A) Perithecium (bar = 100 µm), (B) rigid hairs with inflated tips (bar
= 10 µm). Podospora longicaudata. (C) Mature ascus (bar = 10 µm), D. ascospore (bar = 10 µm).
Podospora ostlingospora. (E) Ascospore (bar = 15 µm). Podospora pauciseta. (F) Mature ascus
(bar = 10 µm). (G) perithecium (bar = 100 µm), (H) ascospore (bar = 10 µm). Podospora
pleiospora. (I) Tubercles on the perithecial neck (bar = 7.5 µm), (J) Mature ascus (bar = 12.5 µm),
(K) immature ascospore (bar = 5 µm), (L) Mature ascospore (bar = 12.5 µm). Podospora
prethopodalis. (M) Mature ascospore (bar = 15µm).
214
The main distinguishing feature of this species is the gelatinous equipment, composed of (1) a
single lash-like apical caudae, (2) a single lash-like basal caudae, usually collapsing, attached to
distal end of the pedicel, and (3) a single lateral caudae attached to the proximal end of the pedicel.
Material not examined in this study.
Podospora dolichopodalis J.H. Mirza & Cain, Can. J. Bot. 47(12): 2018 (1970) [1969]
This species was first described by Mirza & Cain (1969) from material growing in dung collected
in Toronto, Canada. It is a peculiar species, in which the ascospore pedicel possesses a conspicuous
distal swelling, appearing somewhat clavate rather than cylindrical. It was recorded in Minas
Gerais, Southeast Brazil (Lundqvist, 1973; Jahn, 2000). Material not examined in this study.
Podospora immersa (R. Stratton) Cain, Can. J. Bot. 40: 460 (1962)
Basionym: Pleurage immersa R. Stratton, Ohio St. Univ. Bull. 26: 93 (1921)
As in Podospora longicaudata, P. immersa has several small caudae completely enveloping
the head cell in its ascospores. It can be distinguished, however, from the former by the narrower
head cells (30-43µm) and, additionally from P. ostlingospora by the shorter caudae. It was also
recorded in Minas Gerais (Lundqvist, 1973; Jahn, 2000). Material not examined in this study.
Podospora longispora (Bat. & Pontual) N. Lundq., Symb. Bot. Upsal. 20(1): 135 (1972)
Basionym: Sordaria longispora Bat. & Pontual, Bol. Secr. Agric. (Pernambuco) 15: 39 (1948)
After studying the type material labeled as Podospora longispora, collected on horse dung
from Dois Irmãos, Pernambuco, Mirza & Cain (1969) placed it as a synonym of P. appendiculata.
Lundqvist (1972), however, examined the same original material, deposited in IPA Herbarium
(Pernambuco, Brazil) and stated that, despite the poor state of the exsiccati and the similar rigid,
hyaline-tipped hairs, there are some differences, such as the membranaceous, subopaque, brown
peridium (vs. the pseudobombardioid peridium in P. appediculata) and the slightly shorter dark
ascospore cell (24-29 µm). Material not examined in this study.
Podospora sp.
After a visit to Brazil, Richardson (2001) published a brief note about an unidentified species
of Podospora, which resembled the illustrations of P. ostlingospora in Mirza & Cain (1969) (MJR
60/98). The limited material of this large-spored species prevented it from being formally described
in his paper. However, the author provided a brief description: Perithecia globose, partially
immersed, glabrous or with long, brown, flexuous hyphal hairs, and some P. decipiens-type
papillae. Ascospores with head cells 48-54.5 × 21-27 μm; pedicel slightly clavate distally, 22-30 ×
6-7 μm; caudae inconspicuous, either in the ascus or in free spores. Material not examined in this
study.
Acknowledgements
The authors would like to thank the “Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior” (CAPES) and the “Conselho Nacional de Desenvolvimento Científico e Tecnológico”
(CNPq-Ciência sem Fronteiras; INCT-Herbário Virtual da Flora e dos Fungos) for providing PhD
scholarships to the first author. L.C. Maia acknowledges the research fellowship and grants
provided by CNPq (INCT-HVFF, Protax, Sisbiota).
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www.indexfungorum.org 2015.
... Although they are cosmopolitan, the occurrence records of this group of organisms are more frequent from Europe, North America and a number of Neotropical countries (Brummelen 1967;Lundqvist 1972;Eliasson and Lundqvist 1979;Jahn 2000;Doveri 2004;Bezerra et al. 2008;Calaça and Xavier-Santos 2012). Studies involving coprophilous fungi have recently been conducted in Brazil (Richardson 2001b;Bezerra et al. 2008;Viriato 2008;Melo et al. 2011;, 2015a, 2015bCalaça and Xavier-Santos 2012;Calaça et al. 2013Calaça et al. , 2014, but there are still few data about the distribution of this group in the country, and there are a large number of unexplored areas that could provide essential knowledge of their diversity. This study adds new records of genera and species for Brazil and for the Neotropical Region, with taxonomic descriptions, characterization and geographic distribution of these species. ...
... These results, added to the recent findings reported by Melo et al. (2014Melo et al. ( , 2015aMelo et al. ( , 2015b and , increases the number of species of coprophilous fungi and other fungi from Brazil recorded on dung from 210 (Calaça et al. 2014) to 225. ...
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This study presents the first records from three genera Hypocopra, Pyxidiophora, Zopfiella and the second of the rare species, Cercophora coronata (Cailleux) Udagawa and T. Muroi, for Brazil. It also presents the first record of H. stercoraria (Sowerby) Sacc., P. arvernensis (Breton and Faurel) N. Lundq. and Z. latipes (N. Lundq.) Malloch and Cain for the Neotropical region. Specimens were obtained from dung collected in areas of the Brazilian Savannah cultured in moist chambers. Distribution data, taxonomic descriptions and new substrate records for are presented.
... Clavicipitaceae is a soil saprophyte and potential insect pathogen ( Torres and White 2009 ). Lasiosphaeriaceae is commonly found in herbivore dung ( Melo et al. 2015 ). The largest genus in the family Bolbitiaceae, Conocybe, is commonly found in herbivore dung and grasslands ( Amandeep et al. 2015 ;Joshi et al. 2021 ). ...
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When grazed intensively, grasslands can result in biodiversity loss and increased greenhouse gas emissions. However, when grasslands are maintained using agroecological practices, they can create diverse habitats and act as carbon sinks. Carbon sequestration in soil is mediated by the soil microbial community, which is largely affected by influxes of nutrients and the soil arthropod community. One such arthropod, the tunneling dung beetle Onthophagus taurus (Schreber 1759) incorporates animal dung into the soil, influencing both the soil microbial community and nutrient cycling. While dung beetles do affect the soil microbial community, there is a dearth of studies that examine the effect of tunneling dung beetles on soil microbial communities that may be associated with nutrient cycling. This study looks at the effect of the tunneling dung beetle O. taurus on the fungal, bacterial, and archaeal community of grassland soil in an in situ experiment in California’s Central Coast region, using eDNA metabarcoding of the 16S locus for bacteria and archaea and the ITS1 locus for fungi. We find that O. taurus has a clear effect on the soil microbial community and its presence increases the abundance of soil microbes affiliated with degrading plant materials and carbohydrate metabolism.
... These included plant pathogens as Camarosporium spp. 109 (9.7% of the OTUs) and Ustilago striliformis 110 , Entyloma microsporum 111 , Microbotrium silybum 112 , accounting overall for a further 4.4% of the OTUs, and coprophila fungi as Podospora (7.5% of OTUs) and Sporormiella (3.0% of OTUs) 113 , the spores of which are ingested by the grazing herbivores and subsequently germinate in dung, so that they are also used as indices of variation through time of large herbivore populations 114 . All together, these taxa comprised more than 25% of the OTUs. ...
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Diet selection by grazing livestock may affect animal performance as well as the biodiversity of grazed areas. Recent DNA barcoding techniques allow to assess dietary plant composition in faecal samples, which may be additionally integrated by the description of gut microbiota. In this high throughput metabarcoding study, we investigated the diversity of plant, fungal and bacterial taxa in faecal samples of lactating cows of two breeds grazing an Alpine semi-natural grassland during summer. The estimated plant composition of the diet comprised 67 genera and 39 species, which varied remarkably during summer, suggesting a decline of the diet forage value with the advancing of the vegetative season. The fungal community included Neocallimastigomycota gut symbionts, but also Ascomycota and Basidiomycota plant parasite and coprophilous taxa, likely ingested during grazing. The proportion of ingested fungi was remarkably higher than in other studies, and varied during summer, although less than that observed for plants. Some variation related to breed was also detected. The gut bacterial taxa remained stable through the summer but displayed a breed-specific composition. The study provided insights in the reciprocal organisms’ interactions affecting, and being affected by, the foraging behaviour: plants showed a high temporal variation, fungi a smaller one, while bacteria had practically none; conversely, the same kingdoms showed the opposite gradient of variation as respect to the animal host breed, as bacteria revealed to be the group mostly characterized by host-specificity.
... Viriato (2008) identified 10 species of Pilobolus in a Zoological Park in São Paulo, and Melo et al. (2011) recorded 22 species of coprophilous ascomycetes on captive wild herbivore dung in a Zoological Park in Recife, providing illustrations, descriptions and an identification key. Recently, more focused works on the most common genera were conducted, and new species and records have been presented (Melo et al. 2014(Melo et al. , 2015a(Melo et al. , 2015b(Melo et al. , 2017a(Melo et al. , 2017b(Melo et al. , 2019. Calaça et al. (2014) provided a preliminary checklist of coprophilous fungi and Myxomycetes from Brazil, as well as other contributions to the knowledge of the coprophilous mycobiota of the Brazilian Cerrado (Calaça et al. 2013, Calaça & Xavier-Santos 2014, 2016, Calaça et al. 2015. ...
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Taxonomic records of coprophilous fungi from Brazil are revisited. In total, 271 valid species names, including representatives of Ascomycota (187), Basidiomycota (32), Kickxellomycota (2), Mucoromycota (45) and Zoopagomycota (5), are reported from herbivore dung. Identification keys for coprophilous fungi from Brazil are provided, including both recent surveys (2011–2019) and historical literature.
... Niessl, but could be distinguished based on spore size, shape and characteristics of the caudae (Mirza & Cain 1969, Bell 2005, Melo et al. 2015. ...
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In a study on coprophilous fungi, different samples including cow, sheep and horse dung and mouse feces were collected from different locations in West and East Azarbaijan provinces (NW Iran). Isolation of the fungi was done based on moist chamber culture method. Purification of the isolated fungi was done by single spore culture method. Several fungal taxa were obtained. Identification of the isolates at species level was done based on morphological characteristics and data obtained from internal transcribed spacer (ITS) regions of ribosomal DNA sequences. In this paper, five taxa viz. Arthrobotrys conoides, Botryosporium longibrachiatum, Cephaliophora irregularis, Oedocephalum glomerulosum, and Podospora pauciseta, all of them belong to Ascomycota, are reported and described. All these taxa are new records for Iran mycobiota.
... However, species of Lasiosphaeria do not form ascospores with a swollen, pigmented head (Seaver 1912;Taylor et al. 2001;Miller and Huhndorf 2001). Members of Arnium and Podospora generally grow on dung and produce ascospores with gelatinous appendages (Krug and Cain 1972;Lorenzo and Havrylenko 2001;Melo et al 2015). Zopfiella has septate ascospores with a hyaline pedicel which often collapses (Cai et al. 2006). ...
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Fourty-three species of microfungi from bamboo are treated, including one new family, Occultibambusaceae, three new genera, Neoanthostomella, Occultibambusa and Seriascoma, 27 new species, one renamed species and 15 re-described or re-illustrated species, and four designated reference specimens are treated in this paper, the majority of which are saprobic on dead culms. To determine species identification, separate phylogenetical analyses for each group are carried out, based on molecular data from this study and sequences downloaded from GenBank. Morphologically similar species and phylogenetically close taxa are compared and discussed. In addition a list of bambusicolous fungi published since Hyde and colleagues in 2002 is provided.
Thesis
We define copromycodiversity as all the morphological, physiological, functional, and ecological diversity of an assemblage of dung-inhabiting fungi, also known as coprophilous fungi. These fungi are represented by species of the phyla Ascomycota, Basidiomycota, and Mucoromycota in the sensu stricto of the Kingdom. Traditionally, other groups of organisms, such as some species of protists from the phylum Amoebozoa (Myxomycetes) can also be found associated with these substrates, being studied together with fungi, here in sensu lato. This fungal group is responsible for the maintenance of supporting ecosystem services, essential for life as we know it, acting in the cycling of matter and energy via dung decomposition. Despite this, there are still few studies involving these organisms. In this thesis, we seek to contribute to expanding scientific knowledge about dung-inhabiting fungi (sensu lato), whether in terms of their biology, ecology, or taxonomy. We begin by promoting a synthesis of knowledge as well as a scientometric analysis of the first studies in Brazil and in the world, in addition to an updated list of species and their geographic distribution, adding new occurrences. We compile data on the occurrence and applications of the genus Chaetomium, a fimicolous fungal genus, which means that its species are found associated with both dung and decaying plant debris, as well as developed a proposal for a life cycle for dung-inhabiting fungi, contemplating both the coprophilous and fimicolous cycles. We show how differences in the type of digestive tract of the animal that originated the dung can influence the taxonomic diversity and ecological profiles of species of fungi that inhabit this substrate. Furthermore, we evaluated, in a microcosm experiment, the durability, decomposition rate, and nutritional composition of cattle and equine dung as suitable substrates for the development of coprophilous organisms. Finally, we present the experience report on how we are using social media to communicate fungal biology to the public, developing media strategies to publicize different aspects of our research with dung-inhabiting fungi as well as other fungal groups to the general community. It is expected that the data presented here can contribute to the expansion of knowledge about these fungi, reducing the gap in knowledge about biology, ecology, distribution, and productivity, especially in Brazil. In addition, we hope to encourage the training of human resources as well as the development of additional studies that will contribute to an increasingly better understanding of copromycodiversity.
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Mycotaxon is pleased to add a new annotated species distribution list to our 142 previously posted free access fungae. The 22-page "Additions to a checklist of coprophilous fungi and other fungi recorded on dung from Brazil: an overview of a century of research" by Francisco J. Simões Calaça, Vanessa Basilio Tereza, and Solange Xavier-Santos may be downloaded from our website via http://www. mycotaxon. com/mycobiota/index. html
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In 2014 we published the first checklist of fungi sensu lato recorded on dung from Brazil, as a result of an effort to maintain an easily accessible database on the knowledge of the diversity of these fungi. In the last 10 years there has been a significant increase in the number of publications involving this ecological group, and we are presenting an update which summarizes data from a century of research (from 1919 to 2019). We present an annotated list of all species and occurrences added to the Brazilian copromycodiversity, the annual accumulation of records, and occurrence by substrate. There were 302 records of 166 species, 98 of which are new for Brazil, all of them reported from seven Federation states (including two new). Most of the occurrences are reported from the northeast region of the country (Pernambuco and Piauí states), followed by the midwest (Goiás and Federal District) and southern regions (Paraná and Rio Grande do Sul). Amazonas and Piauí are states with new records of these organisms, with one record each. There are now 568 occurrences and 308 species of dung-inhabiting fungi recorded in Brazil.
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Knowledge of the relationships and thus the classification of fungi, has developed rapidly with increasingly widespread use of molecular techniques, over the past 10–15 years, and continues to accelerate. Several genera have been found to be polyphyletic, and their generic concepts have subsequently been emended. New names have thus been introduced for species which are phylogenetically distinct from the type species of particular genera. The ending of the separate naming of morphs of the same species in 2011, has also caused changes in fungal generic names. In order to facilitate access to all important changes, it was desirable to compile these in a single document. The present article provides a list of generic names of Ascomycota (approximately 6500 accepted names published to the end of 2016), including those which are lichen-forming. Notes and summaries of the changes since the last edition of ‘Ainsworth & Bisby’s Dictionary of the Fungi’ in 2008 are provided. The notes include the number of accepted species, classification, type species (with location of the type material), culture availability, life-styles, distribution, and selected publications that have appeared since 2008. This work is intended to provide the foundation for updating the ascomycete component of the “Without prejudice list of generic names of Fungi” published in 2013, which will be developed into a list of protected generic names. This will be subjected to the XIXth International Botanical Congress in Shenzhen in July 2017 agreeing to a modification in the rules relating to protected lists, and scrutiny by procedures determined by the Nomenclature Committee for Fungi (NCF). The previously invalidly published generic names Barriopsis, Collophora (as Collophorina), Cryomyces, Dematiopleospora, Heterospora (as Heterosporicola), Lithophila, Palmomyces (as Palmaria) and Saxomyces are validated, as are two previously invalid family names, Bartaliniaceae and Wiesneriomycetaceae. Four species of Lalaria, which were invalidly published are transferred to Taphrina and validated as new combinations. Catenomycopsis Tibell & Constant. is reduced under Chaenothecopsis Vain., while Dichomera Cooke is reduced under Botryosphaeria Ces. & De Not. (Art. 59).
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The phylogenetic relationships of Lasiosphaeriaceae are complicated in that the family is paraphyletic and includes Sordariaceae and Chaetomiaceae, as well as several polyphyletic genera. This study focuses on the phylogenetic relationships of the coprophilous genera, Anopodium, Apodospora, Arnium, Fimetariella and Zygospermella. They are traditionally circumscribed based on ascospore characters, which have proven homoplasious in other genera within the family. Our results based on LSU nrDNA and ß–tubulin sequences distinguish four lineages of Lasiosphaeriaceae taxa. Anopodium joins the clade of mor- phologically similar, yellow-pigmented species of Cercophora and Lasiosphaeria. Apodospora is monophyletic and joins a larger group of taxa with unclear affinities to each other, while Arnium is polyphyletic being scattered throughout three of the four major clades of Lasiosphaeriaceae. Fimitariella is represented by a single collection and joins the clade containing Cercophora scortea and Podospora appendiculata. Zygospermella shows affinities to the Lasiosphaeris clade. Based on a combination of morpholog- ical and molecular data, Echria stat. nov. is recognized at the genus level for the former Arnium section and two new combinations are proposed: E. gigantospora and E. macrotheca.
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Coprophilous fungi are highly satisfactory for demonstrating the diversity and morphology of a group of related organisms within an ecological system. Representative genera of most major groups of fungi can usually be guaranteed to appear on dung after a period of incubation in a damp chamber. Four keys are presented. They are not exhaustive, since there are far too many species to make it practical to include them all. They do, however, include most genera, and the commoner or well-known species of temperate regions. There are two keys to about 270 coprophilous ascomycetes, a very diverse group which, although not covering all the possible types of reproductive structure found in the class, contains many of the important types. A third key is to about 75 basidiomycetes of dung and associated debris. Key 4 is to representative species of those Zygomycota (phycomycetes) which are characteristic of dung and amongst the first to appear when fresh dung is incubated.
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K.D. (2005). Phylogenetic evaluation and taxonomic revision of Schizothecium based on ribosomal DNA and protein coding genes. Fungal Diversity 19: 1-21. The taxonomy of Schizothecium and Podospora has been a subject of debate. Both of these genera have previously been treated as congeneric due to the lack of distinguishing morphological characters. This study focuses mainly on the phylogenetic relationships of Schizothecium and Podospora, and also a re-evaluation of the taxonomic significance of morphological characters. Multiple gene sequences (partial 28S rDNA, ITS/5.8S rDNA and partial β-tubulin) were analysed using maximum parsimony and Bayesian analyses. In all analyses, Schizothecium species characterised by perithecia adorned with swollen agglutinated hairs or prominent protruding peridial cells, grouped in a strongly supported monophyletic clade. Schizothecium should therefore, be given appropriate generic status and not treated as congeneric to Podospora. Phylogenetic analyses provide good support indicating that ascomatal morphologies are more phylogenetically informative than ascospore characters and host or habitat association. A synopsis of and key to Schizothecium species now recognised in the genus are given. Podospora was found to be a polyphyletic genus, consisting of a group of morphologically heterogeneous and phylogenetically distant species. Preliminary data indicate that a complete revision of Podospora and related genera is necessary.
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The genus Lasiosphaeria recently has been circumscribed more narrowly to include five mor-phospecies united by tomentose ascomata containing yellow centrum pigments. Species boundaries have not been established and phylogenetic relationships have not been clearly defined for these morphospecies. To delimit species boundaries and determine phylogenetic relationships among species, maximum parsimony, maximum likelihood and Bayesian analyses were conducted on sequence data from four nuclear genes, the ribosomal internal transcribed spacer (ITS) region, 28S large subunit (LSU) rDNA, β-tubulin and ribosomal polymerase II subunit 2 (RPB2). Representatives of L. glabrata, L. ovina, L. rugulosa and L. sorbina resolved as four highly supported monophyletic groups in almost all analyses and are recognized as well-defined species employing principles of genealogical concordance. These species delimitations are corroborated further by morphology. Representatives of L. lanuginosa were polyphyletic in almost all analyses. Although molecular analyses revealed that this morphospecies comprises several phylogenetic species, formal taxonomic recognition of these lineages is premature, so L. lanuginosa currently is treated as a morphological species complex. Complete species descriptions, including teleomorph, anamorph and culture characteristics, are given for L. glabrata, L. ovina, L. sorbina and the L. lanuginosa species complex along with detailed discussions of significant morphological characters used in recognizing species. These species are compared to five additional morphospecies that also may belong in the genus.
Article
Twenty-two species of coprophilous ascomycetes were identified from 216 samples of dung from confined paca, tapir, deer, llama, camel and waterbuck collected in the Parque Estadual Dois Irmãos, Recife, Brazil. Members of Ascobolaceae and Chaetomiaceae showed the higher number of occurrences, and Lasiosphaeriaceae the largest number of species present. The genus Saccobolus proved to be the most representative, and Saccobolus saccoboloides was dominant, followed by Saccobolus citrinus, Chaetomium bostrychodes, and Thielavia terrestris. Camel dung showed higher species diversity, and llama dung had higher number of ocurrences. Tapir dung proved unfavorable to the sampling of ascomycetes. The community showed high diversity (H' = 2.54), homogeneous distribution (J = 0.82), and substrate preferences tending to generalism (B = 4.64). The communities in deer and waterbuck dung and in camel and llama dung showed, in pairs, the largest similarity, probably due to the diet similarity. The mycobiota of paca and waterbuck dung showed no similarity, and the mycobiota in tapir dung showed no similarity to any other. The communities studied showed low temporal increase in species composition throughout the year, with high diversity and frequent sampling of rare species, no significant relation between digestive system type and a significant difference between substrate and month of collection.
Article
A key is provided for the 64 recognized species of Podospora. The genus is restricted to species in which the ascospores have an apical germ pore, a basal hyaline cell, and gelatinous appendages. Illustrations are given for 56 species. Detailed descriptions are included for species which are not readily available and the remaining species are briefly described. The following 10 new species found on dung are described and illustrated: P. attoeochaeta from Mexico; P. didyma, Quebec; P. dolichopodalis, Louisiana and Mexico; P. gigantea, Louisiana and Arkansas; P. karachiensis, Pakistan; P. macropodalis, Mexico; P. mexicana, Mexico; P. miniglutinans, Mexico; P. pistilata, Mexico; P. venezuelensis, Venezuela, Mexico, Louisiana, South Pacific. The following new combinations are proposed: P. aloides (Sordaria aloides Fuckel), P. apiculifera (S. apiculifera Speg.), P. argentinensis (S. argentinensis Speg.), P. austro-americana (Hypocopra austro-americana Speg.), P. ellisiana (Pleurage ellisiana Griff.), P. longicollis (Schizothecium longicollis Ames), P. seminuda (Sordaria seminuda Griff.), P. tarvisina (Philocopra tarvisina Sacc.), and P. vesticola (Sphaeria vesticola Berk. & Br.).
Article
Podospora is the valid genus for a group of coprophilous species in which the ascospores have appendages but no gelatinous sheaths. The type species is P. fimicola Cesati. The following new species found on dung are described and illustrated: P. absimilis, Tautira; P. brasiliensis, Brazil; P. castorinospora, Panama Canal Zone; P. comatospora, Venezuela, Tautira, and Moorea; P. curvuloides, Brazil; P. inflatula, Tautira and Brazil; P. nannopodalis, New York; P. ostlingospora, Mexico; P. papilliformis, Venezuela and Mexico; P. prethopodalis, Tautira, Moorea, and Paea. The following are transferred from Pleurage as new combinations in Podospora: P. adelura (Griff.), P. anomala (Griff.), P. arizonensis (Griff.), P. collapsa (Griff.), P. gwynne-vaughaniae (Page), P. heterochaeta (Griff.), P. hyalopilosa (Stratton), P. immersa (Stratton), P. kansensis (Griff.), P. longicaudata (Griff.), P. taenioides (Griff.) and P. unicaudata (C. & M. Moreau). The following are transferred from Sordaria as new combinations in Podospora: P. araneosa (Cain), P. cervina (Cain), P. coronifera (Grove), P. curvispora (Cain), P. dubia (Hansen), P. eminens (Cain), P. fimbriate, (Bayer), P. globosa (Massee & Salmon), P. glutinans (Cain), P. inaequalis (Cain), P. inaequilateralis (Cain), P. kilimandscharica (A. Schmidt), P. leporina (Cain), P. linguiformis (Cain), P. ontariensis (Cain), P. perplexens (Cain), P. pilosa (Mouton), P. piriformis (Bayer), P. tetraspora (Winter). The following are transferred from Philocopra as new combinations in Podospora: P. coeruleotecta (Rehm) and P. millespora (A. Schmidt).