First contribution to the genus Gymnopilus (Agaricales, Strophariaceae) in Paraguay

Abstract

Gymnopilus is characterized by its ferruginous-yellow basidiomata and lamellae, ferruginous spore print, ellipsoidal basidiospores with warty and rough ornamentation, and lacking a germinative pore. Here, novel data on the Gymnopilus species of Paraguay is presented, macro and microscopic morphological characteristics, distribution, and ecology are described, and a taxonomic discussion is provided. Gymnopilus imperialis is recorded in the Alto Paraná Department, G. lepidotus in the Central Department, G. luteofolius in the Cordillera Department, G. peliolepis in the Paraguarí Department, and G. purpureosquamulosus in the Central Department and Boquerón, all as new records for Paraguay. Photographs of the fresh basidiomata and some microscopic structures such as basidia and basidiospores are attached.

Full text

Rodriguésia 72: e00752019. 2021
http://rodriguesia.jbrj.gov.br
DOI: http://dx.doi.org/10.1590/2175-7860202172013
Abstract
Gymnopilus is characterized by its ferruginous-yellow basidiomata and lamellae, ferruginous spore print,
ellipsoidal basidiospores with warty and rough ornamentation, and lacking a germinative pore. Here, novel
data on the Gymnopilus species of Paraguay is presented, macro and microscopic morphological characteristics,
distribution, and ecology are described, and a taxonomic discussion is provided. Gymnopilus imperialis
is recorded in the Alto Paraná Department, G. lepidotus in the Central Department, G. luteofolius in the
Cordillera Department, G. peliolepis in the Paraguarí Department, and G. purpureosquamulosus in the Central
Department and Boquerón, all as new records for Paraguay. Photographs of the fresh basidiomata and some
microscopic structures such as basidia and basidiospores are attached.
Key words: Agaricomycetes, fungal diversity, taxonomy.
Resumen
Gymnopilus se caracteriza por poseer basidiomas y laminillas con tonos amarillo ferrugíneos, esporada
ferruginosa, basidiosporas elipsoidales con ornamentación verrugosa a rugosa, sin poro germinativo. En el
presente trabajo se proporcionan datos novedosos sobre las especies de Gymnopilus de Paraguay, se describen
sus características morfológicas, su distribución, ecología y se proporciona una discusión en torno a su
taxonomía. Se citaron G. imperialis para el Departamento Alto Paraná, G. lepidotus para el Departamento
Central, G. luteofolius para el Departamento Cordillera, G. peliolepis para el Departamento Paraguarí, y
G. purpureosquamulosus para el Departamento Central y Boquerón, y todas como nuevos reportes para el
Paraguay. Además, se ilustran los caracteres microscópicos distintivos. Se anexan fotografías de los basidiomas
en fresco y de algunas estructuras microscópicas como basidios y basidiosporas.
Palabras clave: Agaricomycetes, diversidad de hongos, taxonomía.
Original Paper
First contribution to the genus Gymnopilus (Agaricales, Strophariaceae)
in Paraguay
Michelle Campi1,5,7, Yanine Maubet1,6, Emanuel Grassi 2, Nicolás Niveiro3 & Laura Guzmán-Dávalos4
1 Universidad Nacional de Asunción, Facultad de Ciencias Exactas y Naturales, Lab. Análisis de Recursos Vegetales, área Micología, San Lorenzo, Paraguay.
2 Instituto Misionero de Biodiversidad (IMiBio), Pto. Iguazú, Misiones, Argentina. ORCID: <https://orcid.org/0000-0003-4563-056X>.
3 Universidad Nacional del Nordeste, Inst. Botánica del Nordeste, Corrientes, Argentina. ORCID: <https://orcid.org/0000-0002-3265-7061>.
4 Universidad de Guadalajara, Departamento de Botánica y Zoología, Zapopan, Jalisco, México. ORCID: <https://orcid.org/0000-0002-4390-3678>.
5 ORCID: <https://orcid.org/0000-0001-8809-0159>.
6 ORCID: <https://orcid.org/0000-0002-0322-6238>.
7 Author for correspondence: geraldinecampi@gmail.com
Introduction
The genus Gymnopilus P. Karst. includes
more than 200 saprobic species, mainly lignicolous,
which is why they represent an important
component of the wood mycobiota around the
world (Guzmán-Dávalos 2003; Holec 2005; Kirk et
al. 2008). Based on the pigments it contains and the
non-mycorrhizal habit, Kühner (1984) classified
Gymnopilus together with Galerina Earle in the
Strophariaceae family. Later, Singer (1986) placed
it within Cortinariaceae due to the ornamentation
and lack of germinal pore of the basidiospores.
Currently, according to the results of phylogenetic
analysis based on DNA sequences, Gymnopilus is
not related to any of the two families mentioned
above, since it forms an independent clade called

Campi M et al.
2 de 13
Rodriguésia 72: e00752019. 2021
“Gymnopilae” by Matheny et al. (2006) and is
currently included in the family Hymenogastraceae
(Kirk et al. 2008).
Gymnopilus is characterized by its yellow,
ferruginous or purple basidiomata, yellow to
ferruginous lamellae, central to eccentric stipe,
with a partial cortinoid or fibrillose veil, generally
fugacious, or as a membranous ring, bitter
to farinaceous flesh, ferruginous spore print,
basidiospores ellipsoidal with warty to rugose
ornamentation, without germinative pore and in
most species with dextrinoid walls, presence of
subcapitulated cheilocystidia to capitated and
hyphae with clamp connections (Horak 1989;
Guzmán-Dávalos 2003; Holec 2005). An important
ecological character is the lignicolous habit, being
able to grow in woods in different degrees of
decomposition (Guzmán-Dávalos 2003; Holec
2005) and apparently without having a preference
for the substrate (Singer 1951). However, it has
been observed that some tropical species are
associated with Angiosperms and others with
palms, while in temperate zones they are associated
with conifers.
In South America Gymnopilus has been
recorded by Berkeley & Cooke (1876), Saccardo
(1887), Patouillard & Gaillard (1888), Singer
(1951, 1953b, 1961, 1969, 1975), Singer & Digilio
(1951), Dennis (1953, 1961, 1970), Raithelhuber
(1974, 1980, 1991, 2004), Talice & Talice (1980),
Bononi et al. (1984), Lazo (1984), Garrido (1985,
1988), Horak (1989), Pegler (1988, 1990, 1997),
Putzke (1994), Valenzuela et al. (1994), Sede
& López (1999), Cardona et al. (2005), Cortez
& Coelho (2005), Franco-Molano et al. (2005),
Wright & Wright (2005), Lechner et al. (2006),
Drechsler-Santos et al. (2007), Karstedt & Stürmer
(2008), Wright et al. (2008), Lobato et al. (2010),
Vasco-Palacios & Franco-Molano (2013), Magnago
et al. (2013), Sequeira (2013), Silva-Junior &
Wartchow (2015) and Grassi et al. (2016). These
authors have cited species for Argentina, Bolivia,
Brazil, Chile, Colombia, Ecuador and Uruguay.
Nonetheless, records of this genus in Paraguay
are scarce: Saccardo (1887), who cited Flammula
sapinea var. australis Kalchbr. and Piris da Motta
et al. (2015), who cited G. subtropicus Hesler for
the Paraguarí Department. However, Gymnopilus
presents an important diversity in Paraguay, which
is not yet documented. The aim of this work was to
describe species of Gymnopilus and thus contribute
to the registration and knowledge of agaricoid
mycobiota of Paraguay.
Materials and Methods
Samples were collected in the Departments
Alto Paraná, Boquerón, Central, Cordillera and
Paraguarí-Paraguay, and analyzed in the Análisis
de Recursos Vegetales Laboratory, Mycology lab,
Facultad de Ciencias Exactas y Naturales (FACEN),
of the Universidad Nacional de Asunción, Paraguay.
The macroscopic descriptions were based on fresh
material, according to the guidelines proposed by
Wright & Albertó (2002) and Lodge et al. (2004).
The microscopic characteristics were described
from the material mounted in 5% KOH and
observed in an optical microscope; in addition,
1% floxin, Congo ammoniac red, Melzer reagent
and cresyl blue were used, the last one for the
metachromatic reaction. The following notations
were followed when performing the measurements
of the basidiospores: Q = ratio between the length
and width of the spores, indicated as a range of
variation; n = number of measured spores and N =
number of basidiomata (Niveiro et al. 2012). The
specimens studied were deposited in the herbarium
of the Facutad de Ciencias Exactas y Naturales of
the Universidad Nacional de Asunción, Paraguay
(FACEN).
Results and Discussion
Five species of Gymnopilus were recorded:
G. imperialis in the Alto Paraná Department, G.
lepidotus in the Central Department, G. luteofolius
in the Cordillera Department, G. peliopelis in the
Paraguarí Department and G. purpureosquamulosus
in the Central and Boquerón Departments, all as
new records for Paraguay.
Gymnopilus imperialis (Speg.) Singer, Lilloa 22:
561 (1951) [1949].
≡ Pholiota imperialis Speg., Boletín de la
Academia Nacional de Ciencias en Córdoba 11(4):
416 (1889). Fig. 1
Cespitose to scattered basidiomata. Pileus
8–12 cm in diameter, ovoid when immature, convex,
plane-convex to flat; margin entire, wavy, revolute
in maturity; surface wet to dry, ochraceus orange,
finely velvety when young, turning ferruginous
orange, radially fibrillose in mature specimens,
not hygrophane. Lamellae adnate to sinuated,
ventricose, up to 0.8 cm wide, subdecurrent to
decurrent, yellow when young, tobacco color to
ferruginous when old, margin entire, concolorous,
lamelullae of two lengths. Ferruginous spore
print. Context ferruginous orange, does not

Gymnopilus in Paraguay 3 de 13
Rodriguésia 72: e00752019. 2021
change to the touch. Characteristic smell of
tobacco, taste not tested. Stipe 6–12 × 0.8–2 cm,
cylindrical, widened towards the base, central,
hollow, flaring, sulphurous yellow when young,
reddish brown when mature, fibrous to fibrillose.
Membranous partial veil, sulfurous yellow, later
as an apical, evanescent membranous ring that
leaves membranous remains to rust-brown fibrils
in the stipe. Abundant basal mycelium, yellowish.
Spore print dark brown ferruginous.
Basidiospores (6.4–)8.5–9 × 4.4–6.8 µm,
Q = 1.26–1.44, Qx = 1.35, n = 30, N = 2, widely
ellipsoid to ellipsoid, apex rounded to acute,
strongly verrucose, warts large, without plage,
with or without suprahilar depression, golden
to ferruginous, dextrinoid, not metachromatic.
Basidia (25–)28–35 × 7–8 µm, clavate, tetrasporic,
sterigmata 4.5–6 μm long, amber to ferruginous.
Pleurocystidia absent. Cheilocystidia 27.5–38.5
(–40) × 5.5–10 µm, clavate to narrowly utriform,
subcapitated to capitated. Subhymenium cellular.
Hymenophoral trama subregular, hyphae 4.5–9
(–11.5) μm in diam., thin-walled, yellowish. Pileus
trama interwoven, hyphae of 5.5–8.5(–13.5) μm
diam., thin-walled, yellowish. Pileipellis a cutis,
prostrate hyphae, some erect to suberect, amber
to brownish, septate, thin-walled. Caulocystidia
not observed. Clamp connections present in the
hyphae of the hymenophoral trama, pileus trama
and stipe.
It was found in grassland near plantations
of Pinus elliotti, during autumn.
Figure 1 – a-f. Gymnopilus imperialis – a, b, c. basidiomata when fresh (b. inmature basidioma); d. cheilocystidia;
e. basidiospores in cotton blue; f. basidiospores in KOH. (leg. M. Campi 027).
5 cm 5 cm
2 cm
10 µm 7 µm 7 µm
a
b
c
ef
d

Campi M et al.
4 de 13
Rodriguésia 72: e00752019. 2021
The type specimen is from Brazil, also
recorded from Argentina, Costa Rica and Jamaica
(Spegazzini 1889; Hesler 1969; Guzmán-Dávalos
et al. 2003). In Brazil it has been cited for the states
of São Paulo and Paraná (Spegazzini 1889; Pegler
1997; Capelari et al. 2015; BFG 2018).
Examined material: PARAGUAY. ALTO PARANÁ:
Refugio Biológico Tatí Yupí, 25°22’22.6”S,
54°35’50.5”W, 10.IV.2015, M. Campi 003346 (FACEN).
This species is grouped within the spectabilis-
imperialis clade as the basal species, which
also includes G. junonius (Fr.) P.D. Orton, G.
pampeanus (Speg.) Singer and G. spectabilis (Fr.)
Singer, this clade includes robust basidiomata, with
thick membranous ring and fibrillose to slightly
squamulosus, dark ferruginous colour pileus
(Guzmán-Dávalos et al. 2003). The species of this
clade are very similar to each other, and in many
cases have been considered as synonyms or varieties
of the same taxon (Guzmán-Dávalos et al. 2003).
Gymnopilus pampeanus is a related species,
and is differentiated from G. imperialis by its
cylindrical stipe and adnexed to sinuate-adnate
lamellae in the first one, besides G. pampeanus
appears to be closely associated with Eucalyptus
forests (Pegler 1983, 1997).
Another closely related species is G. rugulosus
R.Valenz, Guzmán & J. Castillo. This species is
known for the tropical and subtropical regions of
the northern hemisphere, shares characters such as
robust basidiomata with tuberculated basidiospores;
however, it differs from G. imperialis by the
presence of lageniform, ventricular, non-capitated
or subcapitulated pleurocystidia of 21–38 × 5–10
μm (Guzmán-Dávalos & Ovrebo 2001; Cardona
et al. 2005).
Gymnopilus imperialis was originally
described for the south of Brazil (Spegazzini,
1889), and considering the main morphological
characters like robust basidiomata, pileus greater
than 10 cm in diameter, claviform stipe widened
towards the base, membranous ring, basidiospores
subglobose to ellipsoidal, tuberculated, and
absence of pleurocystidia to be consistent with
those observed in the Paraguayan specimen, the
material analyzed from Paraguay is identified as
G. imperialis.
Gymnopilus lepidotus Hesler, Mycologia Memoirs
3: 40 (1969). Fig. 2
Basidiomata cespitose. Pileus 3,5–10 cm
diam., hemispheric to convex, margin entire when
young, plane-concave with revolute margin at
maturity; wet to dry surface, ferruginous chestnut
with vinous tones, becoming clear towards the
margin, covered with chestnut scales, erect,
abundant in the center, scattered towards the
margin. Lamellae subdecurrent, crowded, up to
0.4 cm wide, reddish brown in fresh, ferruginous
in dry, margin entire, concolorous, lamelullae
of two lengths. Context thin, up to 0.1 cm thick,
ferruginous yellow in dry samples, darkens to
reddish brown with contact in fresh samples.
Smell and taste not tested. Stipe 2–7 cm × 0,4–1
cm, central, cylindrical, widens slightly towards
the base, up to 1 cm in diameter, hollow, beige to
light brown, fibrillose, striated, reddish to the touch.
Partial veil not observed. Basal mycelium whitish
to cream, abundant. A reddish pigment dissolves
from the pileus when mounted in KOH. Spore print
yellow ferruginous to orange ferruginous.
Basidiospores 6–7.5(–8.5) × (4–)4.5–5 µm,
Q = 1.5–1.53, Qx = 1.51, n = 30, N = 2, ellipsoid,
amygdaliform, with rounded to subacute apex,
without plage, with or without suprahilar depression,
thin-walled, warty, warts medium, golden to ocher,
dextrinoid, not metachromatic. Basidia 19–24 ×
6–7.5 µm, claviform, bi to tetrasporic, sterigmata
4.3-4.5 μm in length, hyaline or with amber-colored
content. Pleurocystidia of pseudocystidia type,
21.5–27.5(–43.5) × 6–7.5(–10) µm, utriform to
subfusiform, with amber content. Cheilocystidia
(15–)18–23 × (6–)8–8.5 µm, lageniform, with
obtuse to subcapitate apex, hyaline or with amber
content. Subhymenium cellular. Hymenophoral
trama subregular, hyphae of 2–4 μm diam., up to
18.5–25.5 μm diam., thin-walled, hyaline or a few
with amber content. Pileus trama radial, hyphae of
2.5–4 μm in diam., thin-walled, hyaline or a few
with amber color. Pileipellis cutis to trichodermis
in the scales, hyphae of 4.5–13 μm in diam.,
orange to yellowish, prostrate, suberect to erect.
Caulocystidia 29.5–32 × 5–5.5 μm, cylindrical,
thin-walled, scarce in the apical portion, absent
in the rest of the stipe. Clamp connections present
in the hyphae of the pileus trama, hymenophoral
trama and stipe.
It was found on decomposing trunk, in urban
area.
The type is from Florida, United States
America (Hesler 1969; Guzmán-Dávalos 2003).
In addition it was recorded from Argentina for
Misiones Province (Wright & Wright 2005;
Lechner et al. 2006; Wright et al. 2008; Grassi et
al. 2016), from Colombia (Franco-Molano et al.
2005; Vasco-Palacios & Franco-Molano 2013) and

Gymnopilus in Paraguay 5 de 13
Rodriguésia 72: e00752019. 2021
from Mexico for the states of Jalisco and Veracruz
(Guzmán-Dávalos 1996).
Examined material: PARAGUAY. CENTRAL: San
Lorenzo city, Universidad Nacional de Asunción
Campus, 25°20’5,51”S, 57°30’56,95”W, 10.II.2017,
Y. Maubet 003783 (FACEN).
Gymnopilus lepidotus is characterized by
the brownish-colored pileus with small erect or
adherent scales, pleurocystidia and basidioles
with granular, golden content, usually very
abundant, cheilocystidia and pleurocystidia not
capitated to subcapitated and radial pileus trama
(Guzmán-Dávalos 1996, 2003).
The size of the pileus is a character that
has been discussed: Hesler (1969) and Wright et
al. (2008) described the pileus as small, 4–8 mm
in diam.; later Guzmán-Dávalos (1996, 2003),
after reviewing the samples from Florida, stated
that the measurements obtained by Hesler (1969)
maybe mistaken because, even when dry, the
samples reviewed by Hesler presented a bigger
pileus, from 6–18(–26) mm in diam. In addition,
some samples from Mexico reached up to 70
mm in diam. As for the pleurocystidia, Hesler
(1969) mentioned that they are hyaline, rare
and inconspicuous; abundant clavate basidioles
with brown content. Guzmán-Dávalos (1996)
interprets these basidioles as pleurocystidia of
the pseudocystidia type, which coincides with the
samples described by Wright et al. (2008), who
mentioned subpyriform, clavate to subutriform
pleurocystidia, 20–22 × 5–6 μm, with ferruginous
chestnut content.
Gymnopilus dilepis (Berk. & Broome)
Singer is a very similar species to G. lepidotus,
and probably are synonymous (Guzmán-
Dávalos 2003). However, G. dilepis has a
paleotropical distribution, whereas G. lepidotus
is an exclusively neotropical species (Guzmán-
Dávalos 2003).
The presence of the cheilocystidia,
caulocystidia, pleurocystidia of the pseudocystidia
type with amber content in the samples from
Paraguay are consistent with the microscopic
Figure 2 – a-f. Gymnopilus lepidotus – a, b. basidiomata (a. basidiomata when fresh; b. detail of the pileus showing
scales); c. basidiospores; d. pleurocystidia of pseudocystidia type; e. cheilocystidia; f. caulocystidia. (leg. Y. Maubet 040).
8 cm
5 cm
10 µm
25 µm 25 µm 20 µm
a
b
c
ef
d

Campi M et al.
6 de 13
Rodriguésia 72: e00752019. 2021
descriptions of Guzmán-Dávalos (1996, 2003).
However, the size of the basidioma in the
Paraguayan samples are significantly greater
than those cited in the literature. The basidiomata
collected here were found saturated with rain
water, which may have influenced the color
of the pileus and the scales. Given that these
characters are controversial, we do not believe
it to be enough to discard the species, thus we
conclude the specimen studied corresponds to
G. lepidotus, and this therefore the first record
in Paraguay.
Gymnopilus luteofolius (Peck) Singer, Lilloa
22: 260 (1951) [1949].
≡ Agaricus luteofolius Peck, Annual Report on
the New York State Museum of Natural History
27: 94 (1875).
Figure 3 – a-h. Gymnopilus luteofolius – a. basidiomata; b. basidiospores; c. hymenophoral trama; d1. basidium; d2.
pleurocystidum; d3. basidiole; e. pileocystidium; f. pleurocystidium with rostrate apex; g. cheilocystidia; h. basidia.
(leg. M. Campi 128).
0,5 cm
10 µm
20 µm
15 µm
6 µm
6 µm6 µm 5 µm
a
b
c
efgh
d

Gymnopilus in Paraguay 7 de 13
Rodriguésia 72: e00752019. 2021
≡ Pholiota luteofolius (Peck) Sacc., Sylloge
Fungorum 5: 756 (1887). Fig. 3
Basidiomata cespitose. Pileus 1.1–2.3 cm
in diam., hemispheric to plane-convex; margin
entire to rimose when fresh, involute when young,
appendiculate when mature, dry surface, whitish
to cream color, covered by fibrillose scales, erect,
suberect to prostrate, reddish to purple, permanent,
distributed more or less evenly over the surface
of the pileus, which in dry turns dark brown on
a yellowish to ocher surface. Lamellae close,
ventricous, up to 0.3 cm wide, with a decurrent
tooth, cream to yellowish in fresh, chestnut golden
when dry. Margin entire, concolorous, lamelullae
of two lengths. Context whitish in fresh, yellowish
in dry. Smell and taste not tested. Stipe 2,3–3,5 ×
0,5–0,8 cm, central, cylindrical, uniform to slightly
widened towards the base, hollow, flexuous,
fibrillose to squamous, scales brown, insert to
the substrate. Partial veil remains as yellowish
membranous remains at the top of the stipe,
evanescent. Spore print ferruginous chestnut.
Basidiospores de 7.3–9.7 × 5–7.2 µm, Q
= 1.4–1.5, Qx = 1.45, n = 30, N = 2, ellipsoidal,
amygdaliform, without plage, with suprahilar
depression, thin-walled, warty, warts medium, golden
to reddish brown, not dextrinoid, not metachromatic.
Basidia 21.5–27.2 × 6.9–9.4 µm, claviform, bi and
tetrasporic, 3.2–4.7 μm long sterigmata, present in
the margin and the edge of the lamellae, hyaline.
Basidioles 16.2–28.3 × 5.9–7.6 μm, thin-walled,
hyaline. Pleurocystidia 24–35 × 6.3–8.3 μm,
fusiform with rostrate apex, thin-walled, hyaline.
Cheilocystidia of 21–34.6 × 4.2–7.6 μm, lageniform,
apex obtuse to subcapitated, hyaline. Subhymedium
ramose inflated. Hymenophoral trama subregular,
hyphae of 2.6–7.5 μm in diam., thin-walled. Pileus
trama interwoven, hyphae of 12.6–24.8 μm in diam.,
thin-walled, loosely arranged, hyaline, towards the
surface of the pileus the hyphae present a refringent
content, 3.9–8.8 μm. Pilleipelis a cutis to a trichoderm
in the scales, with hyphae of 7.5–18.5 μm in diam.,
hyaline to chestnut with embedded pigment, erect
to form scales, thin-walled, sometimes thickened.
Pileocystidia 46–97 × 14–24 μm, pyriform to
fusiform, occasional, septate, thin-walled, hyaline.
Caulocystidia not observed. Clamp connections
present in the hyphae of the pileipellis, pileus trama,
hymenophoral trama and stipe.
It was found in rotting trunk, in urban area.
It's distributed in Argentina and some states of
the United States of America (Singer 1953b; Hesler
1969).
Examined material: PARAGUAY. CORDILLERA:
Atyra City, Los Agüero country house, 25°20’45.2”S,
57°11’19.0”W, 12.XI.2016, M. Campi 128 (FACEN).
This species is characterized by a dark reddish
to reddish brown colour on the surface of the pileus
when young, turning pinkish red or yellowish red
to yellowish; red to vinous context that shifts to
yellowish, and the clavate to ventricose caulocystidia
disposed in clusters, in addition the species has
fusoid to ventricose pleurocystidia and ventricose
cheilocystidia with capitate to non-capitate apex
(Hesler 1969). The studied samples do not present
dextrinoid basidiospores, which differs from the
observations of Hesler (1969). Singer (1969) noted
that the dextrinoid reaction could be lost in herbarium
samples and it is very difficult to observe. Because
the fresh material was not studied for the Paraguayan
material, it is inconclusive whether or not the
basidiospores are dextrinoid.
Gymnopilus peliolepis (Speg.) Singer, Lilloa 22:
561 (1951) [1949].
≡ Pholiota peliolepis Speg., Boletín de la Academia
Nacional de Ciencias en Córdoba 23(3–4): 394
(1919) [1949]. Fig. 4
Basidiomata cespitose. Pileus of 2.5–7.5
cm in diam., convex to plane-convex, margin
entire, involute; surface yellowish chestnut, wet
to dry, covered with erect to flattened scales,
pyramidal, purple, evenly distributed when young,
diminish towards the margins until disappearing
in mature samples, not hygrophanous. Lamellae
close, ventricose, up to 0.4 cm wide, decurrent to
subdecurrent, yellow in fresh, with uneven color,
ferruginous orange, with purple shades on the edge
when dry, margin entire, lamellulae of three lengths.
Context thin, up to 0.8 cm thick, whitish, does not
change color with touch. Smell and taste not tested.
Stipe 2–3 × 0.2–0.6 cm, central, cylindrical, widens
slightly towards the base, up to 0.66 cm diam.,
flexuous, fibrillose, hollow, light yellow to light
brown, stains with ferruginous color towards the
base. Partial veil forms a fibrillose ring in the upper
part of the stipe, fleeting. Basal mycelium cottony,
whitish, adhering to the substrate. Spore print
ferruginous chestnut.
Basidiospores 7.8–9.2(–9.8) × 4.9–6.0 µm,
Q = 1.56–1.64, Qx = 1.60, n = 30, N = 2, ellipsoid,
with rounded to subacute apex, thin-walled, with or
without plage, with or without suprahilar depression,
verrucose, warts medium, ferruginous golden,
dextrinoid, non-metachromatic. Basidia 14.5–18
× 5–6 μm, clavate, tetrasporic, sterigmata 3.5–4

Campi M et al.
8 de 13
Rodriguésia 72: e00752019. 2021
μm long, hyaline. Cheilocystidia 11–28 × 4.5–8.5
μm, lageniform, capitate, hyaline or with chestnut
content. Subhymenium cellular. Hymenophoral
trama subregular, hyphae of 3–12.5(–15) μm, hyaline,
thin-walled. Pileus trama composed of intertwined
hyphae of 3–14.5 μm in diameter, hyaline, thin-
walled. Thrombopleurous hyphae of 4.5–9.5 μm in
diam., with yellowish refringent content, abundant.
Pileipellis a cutis to trichoderm in the scales, with
hyphae of 6.5–11.5 (–13.5) μm diam., prostrate,
suberect to erect, golden to brown. Pileocystidia
not observed. Caulocystidia not observed. Clamp
connections present in the hyphae of the pileus
trama, hymenophore trama and stipe when mounted
in KOH. A yellowish pigment dissolves.
It was found on decaying trunk of the palm
native to south America, Acrocomia aculeata, during
summer.
It's known for Argentina, Brazil and Florida,
USA according to Hesler (1969), Farr (1973),
Raithelhuber (1988, 1991, 2000, 2004), Singer (1951,
1953a) and Singer & Digilio (1951).
Figure 4 – a-f. Gymnopilus peliolepis – a, b, c. basidiomata (b. pileus in lateral view showing stipe insertion; c. pileus
showing surface configuration); d. basidiospores in KOH; e. basidiospores in cotton blue showing ornamentation; f.
cheilocystitum. (leg. C. Mancuello 013).
5 cm
2 cm
2 cm
10 µm10 µm 7 µm
a
b
c
efd

Gymnopilus in Paraguay 9 de 13
Rodriguésia 72: e00752019. 2021
Examined material: PARAGUAY. PARAGUARÍ:
Quiindy city, 25°54’26.9”S, 57°15’56.1”W, 11.XII.2016,
C. Mancuello 004314 (FACEN).
Gymnopilus peliolepis is characterized
by its whitish-yellowish pileus, covered by
fibrillose scales from bright red to dark purple,
basidiospores with small to medium warts, absence
of pleurocystidia and caulocystidia and being
associated with decomposed hardwood (Singer
1951).
A morphologically related species is G.
purpureosquamulosus, which Høiland (1998)
proposes to differentiate specimens collected in
Africa and Asia. Subsequently, Guzmán-Dávalos
et al. (2008) recorded this species in Central
America (Panama) and Europe. Previously it had
been determined as G. palmicola Murrill or as G.
peliolepis; thus G. peliolepis would currently be
restricted to neotropical regions. Høiland (1998)
differentiates G. purpureosquamulosus by its
slightly thinner basidiospores (8.25 × 5.2 μm)
and ventricose cheilocystidia with prostrate apex,
rounded to globose, but not markedly capitated,
as seen in G. peliolepis. Gymnopilus palmicola is
differentiated by its larger basidiospores (8–12 ×
5.6–7.2 μm) with larger warts (Guzmán-Dávalos
et al. 2008).
From the type specimen of Brazil, Guzmán-
Dávalos et al. (2008), described two types of
basidiospore shape: from the spore print of 6–8 ×
4.4–5.2 µm, ellipsoidal, with medium warts, and
from the basidioma, 6.4–7.2 × 4–4.8 um, ellipsoidal
to oblong, with very small warts, almost asperulated.
Singer (1951) and Hesler (1969) described verrucose
basidiospores for this species. The basidiospores of
the studied material are 7.2–9.2 × 4.9–6 µm, which
is somewhat wider than those described by Guzmán-
Dávalos et al. (2008). Singer (1951) described
spores with verrucose ornamentation, which is
consistent with the basidiospores presented.
An important character to consider of G.
aculeatus (Bres. & Roum.) Singer is the habit.
Gymnopilus aculeatus has been found on dead
trunks of palms and orchids (Singer 1951). The
same is described for G. palmicola, which is
always related to palms. On the other hand, G.
purpureosquamulosus grows both on decaying
wood (Høiland 1998; Guzmán-Dávalos et al.
2008) and on living palms (Guzmán-Dávalos et al.
2008), and G. peliolepis is restricted to decaying
wood (Singer 1951). However, more studies are
necessary to determine the substrate specificity of
these species.
Gymnopilus purpureosquamulosus Høil.,
Mycotaxon 69: 82 (1998). Fig. 5
Basidiomata cespitose. Pileus of 2–9 cm
diam., hemispherical, convex to plano-convex at
maturity, margin entire to crenate when young,
smooth to wavy; wet to dry surface, bright yellow
when young, ferruginous chestnut when mature,
covered with erect, triangular, permanent scales,
evenly distributed or grouped in the center and
diminishing towards the margins, in dry samples
the margin scales become appressed, reddish,
ferruginous chestnut to violet when fresh, purple
when dry. Lamellae close, ventricose, up to 0.5 cm
wide, sinuated to shortly decurrent, concolour to the
surface of the pileus in fresh, orange ferruginous
when dry, margin entire, wavy, concolor, lamelullae
of three lengths. Context yellowish up to 0.4 cm
thick. Smell inappreciable, taste not bitter. Stipe
2–7 × 0.4–1 cm, central, cylindrical, widens slightly
towards the base, hollow, flexuous, fibrillose,
whitish to yellowish when fresh, stains to reddish
brown to the touch. Partial veil leaves fibrillose
remnants of ferruginous tonalities in the upper
part of the stipe, fleeting. Basal mycelium whitish,
abundant. Spore print ferruginous.
Basidiospores (5.9–)6.5–8 × 4–5.5 µm; Q
= 1.46–1.51, Qx = 1.48, n = 30, N = 3, ellipsoid,
amygdaliform, with subacute apex, without plage,
with or without suprahilar depression, thin-walled,
finely verrucose, warts small, golden to reddish
brown, dextrinoid, not metachromatic. Basidia
20–30 × 6–9 μm, clavate, tetrasporic, sterigmata
3–5.5 μm long, hyaline or with yellow-brown
cytoplasmic content. Basidiole (16.5–)17.5–35
(–40) × (4–)5–9.5 μm, thin-walled, hyaline or
mostly with golden-brown content. Pleurocystidia
of the pseudocystidia type of 16.5–28(–35) ×
5–9.5 μm, μm, fusiform, hyaline or with uniform
or irregular orange or golden brown content.
Cheilocystidia (12–)16–27.5 × 5–9 μm, lageniform
with obtuse to subcapitated apex, hyaline or with
golden brown coloration. Subhymenium ramose-
inflated. Hymenophoral trama interwoven, hyphae
5–14(–16) μm in diam., thin-walled. Pileus trama
interwoven, with hyphae of (4–)5–15(–17.5) μm
diam., thin-walled, widening towards the superficial
layers. Pileipellis a cutis to trichoderm in the scales,
with hyphae of 4.5–14.5(–17.5) μm in diam.,
prostrate to suberect or erect when they form the
scales, thickened walls, golden brown, with pigment
embedded in its walls. Pileocystidia not observed.
Caulocystidia (38–)47–74.5 × 5.5–14.5 (–15.5) μm,
cylindrical, in erect to recurved tufts, towards the tip

Campi M et al.
10 de 13
Rodriguésia 72: e00752019. 2021
of the stipe, scarce to absent in some samples. Clamp
connections present in the hyphae of the pileipellis,
pileus trama, hymenophoral trama and stipe. When
mounting in KOH a yellowish pigment dissolves.
It was found in decaying wood or trunks,
urban area, found during spring and summer.
The species have pantropical distribution.
Zimbabwe (Høiland 1998), Nigeria, Italy,
Switerland, Panama (Guzmán-Dávalos et al. 2003),
India (Acharya et al. 2017) and Brazil (Neves et
al. 2013).
Examined material: PARAGUAY. CENTRAL: San
Lorenzo City, Universidad Nacional de Asunción
University Campus, 25°20’11,62”S, 57°30’51,98”W,
2.XI.2016, M. Campi 003774 (FACEN). BOQUERON
DEPARTMENT, Montanía country house, 21°57’30.8”S,
60°11’15.6”W, 6.I.2016, Y. Maubet 026 (FACEN).
Gymnopilus purpureosquamulosus is
characterized by having the pileus covered with
pointed and erect scales in the center, adpressed
towards the margin, reddish, lilac, reddish brown
when fresh, and purple when dry, ellipsoidal
to oblong basidiospores, with subacute apex,
the cheilocystidia of varied forms, lageniform,
utriform, cylindrical to fusiform, with claviform,
rounded to globose apex. (Høiland 1998; Guzmán-
Dávalos et al. 2008; Acharya et al. 2017).
Høiland (1998) indicates that G. peliolepis
(Speg.) is similar to G. purpureosquamulosus;
however, it differs in that the former has smaller
basidiospores and less capitated cheilocystidia.
Another morphologically similar species is G.
dilepis, although it has smaller basidiospores than
those found in G. purpureosquamulosus (Høiland
1998). In the description of the type specimen, the
scales are adpressed in the center, while Guzmán-
Dávalos et al. (2008) mention that the scales are
erect in the center and appressed towards the
margin.
There is some confusion between G.
purpureosquamulosus and G. chrysopellus Singer
& Digilio (1951). Gymnopilus chrysopellus was
described by Berkeley & Curtis (1869) as Agaricus
(Flammula) chrysopellus from Cuba. These authors
Figure 5 – a-f. Gymnopilus purpureosquamulosus – a, b, c. basidiomata (c. pileus showing lamellae); d. basidiospores
in KOH; e. basidiospores in cotton blue; f. cheilocystidia. (leg. M. Campi 121).
a
b
c
ef
d
2 cm 2 cm
5 cm
10 µm
6 µm 6 µm

Gymnopilus in Paraguay 11 de 13
Rodriguésia 72: e00752019. 2021
describe “Pileo umbilicato adpresse tomentoso,
quandoque depresso margine subsulcato…”.
Subsequently, Murrill (1913) proposes the
combination of this species to Gymnopilus, adding
the characters of the spores. However, neither of the
two mentioned authors described the coloration of
the pileus and the scales. Singer & Digilio (1951)
describe in detail G. chrysopellus based on material
from Argentina, Brazil, Peru, Martinique, Guyana,
Cuba and USA, noting that it is a common species
throughout tropical and subtropical America. These
authors provide a description with characters
similar to those described by Høiland (1998)
for G. purpureosquamulosus. Singer & Digilio
(1951) describe the pileus surface as having a
“yellow background, with erect scales, concolorous
with the surface, gradually becoming deep
orange to ferrugineus.” Moreover, Hesler (1969)
described G. chrysopellus with a golden yellow
surface, tomentose, sometime with appressed
fibrils. Høiland (1998) when describing a G.
purpureosquamulosus does not compare it with the
Singer & Digilio descriptions of G. chrysopellus.
Considering this, we could currently restrict G.
chrysopellus to the specimens with the pileus
surface fibrillose, golden yellow, without orange
ferruginous scales, leaving the latter as a distinctive
character for G. purpureosquamulosus. To know
the identity of the specimens studied by Singer
& Digilio (1951), is necessary to re-study them,
in order to know with certainty the distribution of
these two conflictive species. The morphological
differences between the material studied in
Paraguay and that recorded in the bibliography of
the type specimen and additional materials of G.
purpureosquamulosus are: (a) erect scales in the
center agreeing with the description of Guzmán et
al. (2008), but differing from the adpressed scales
described by Høiland (1998) for the type specimen,
and (b) hyphae of the pileipellis with pigment
embedded in bands cited by Høiland (1998) and
Guzmán et al. (2008); however, Acharya et al.
(2017) did not describe this character for Indian
materials. Based on the characters presented by the
cited authors, we conclude that the material studied
corresponds to G. purpureosquamulosus, cited for
the first time in Paraguay.
Conclusion
Spegazzini (1919) recorded in Paraguay two
species as Flammula picrea (Pers.) P. Kumm. and
Flammula sapinea (Fr.) P. Kumm. Later, Flecha
& Niveiro (2019) suggested that this two records
were synonyms of Gymnopilus based on the data
available in Mycobank and Index Fungorum
Databases; however, they did not analysed the
samples from Spegazzini to confirm that they
belong to the genus, hence we can not confirm the
presence of Gymnopilus in the previous records
made by Spegazzini in Paraguay. Flecha & Niveiro
(2019) also mention the species Gymnopilus
earlei Murrill recorded by Gullón (2011), which
represents a photographic record since there is not
a physical record of the sample, therefor this is also
an unconfirmed record. After a long hiatus in the
study of Funga in general, and of Gymnopilus in
particular for Paraguay, this study represent the first
contribution towards the description of Gymnopilus
in the country, nevertheless a broader sampling
and the inclusion of molecular data are necessary
to complement the knowledge of Gymnopilus in
Paraguay and in the region.
Acknowledgments
The authors thank Lic. Jissel Armoa and MSc.
Claudia Mancuello, for the help with the collection
of the materials under study; and Prof. Andrea
Weiler, for the collection of the Chaco samples.
And a special thanks to Patricia Kaishian, for the
meticulous revision of the manuscript’s English
and for the valuable contribution to the writing.
References
Acharya K, Paloi S, Dutta AK, Sikder R & Saha T
(2017) Gymnopilus purpureosquamulosus Høil.
(Agaricales, Basidiomycota): a new distributional
record from India. Check List 13: 1-6.
Berkeley MJ & Cooke MC (1876) The fungi of Brazil,
including those collected by J.W.H. Trail, Esq.,
M.A., in 1874. The Journal of the Linnean Society
Botany 15: 363-398.
Berkeley MJ & Curtis MA (1869) Fungi Cubenses
(Hymenomycetes). Journal of the Linnean Society
10: 280-392.
BFG - The Brazil Flora Group (2018) Brazilian Flora
2020: innovation and collaboration to meet Target
1 of the Global Strategy for Plant Conservation
(GSPC). Rodriguésia 69: 1513-1527.
Bononi VL, Mucci ESF, Yokomizo NKS & Guzmán
G (1984) Agaricales (Basidiomycetes) do Parque
Estadual de Campos do Jordão, SP, Brasil. Rickia
11: 85-89.
Capelari M, Cortez VG, Neves MA, Baseia IG,
Wartchow F, Menolli JN, Karstedt F, Oliveira JJS
& Urrea-Valencia S (2015) Agaricales in Lista de
Espécies da Flora do Brasil. Instituto de Pesquisas
Jardim Botânico do Rio de Janeiro. Available at

Campi M et al.
12 de 13
Rodriguésia 72: e00752019. 2021
<http://oradobrasil.jbrj.gov.br/jabot/oradobrasil/
FB125624>. Access on 05 August 2018.
Cardona B, Saldarriaga Y & Guzmán-Dávalos L
(2005) Primer registro de Gymnopilus rugulosus
(Agaricales, Cortinariaceae) de Colombia. Revista
Mexicana de Micología 21: 55-58.
Cortez VG & Coelho G (2005) Additions to the
mycobiota (Agaricales, Basidiomycetes) of Rio
Grande do Sul, Brazil. Iheringia, Série Botanica
60: 69-75.
Dennis RWG (1953) Les Agaricales de l’Ile de la Trinité.
Rhodosporae-Ochrosporae. Bulletin de la Société
Mycologique de France 69: 145-198.
Dennis RWG (1961) Fungi Venezuelani: IV Agaricales.
Kew Bulletin 15: 67-156.
Dennis RWG (1970) Fungus ora of Venezuela and
adjacent countries. Kew Bulletin Additional Series
III, Cramer, Lehre. Pp. 1-584.
Drechsler-Santos ER, Pastorini LH & Putzke J (2007)
Primeiro relato de fungos Agaricales em fragmento
de mata nativa em Frederico Westphalen. Revista
Brasileira de Biociências 5: 471-473.
Farr ML (1973) An annotated list of Spegazzini’s fungus
taxa. Bibliotheca Mycologica 35: 1-1620.
Flecha Rivas AMI & Niveiro N (2019) Checklist of
agaricoid fungi from Paraguay. 133: 729-729.
Franco-Molano AE, Vasco-Palacios AM, López-
Quintero CA & Boekhout T (2005) Macrohongos
de la región del Medio Caquetá-Colombia. Guía
de campo. Universidad de Antioquia, Medellín.
Pp. 1-211.
Garrido N (1985) Index agaricalium chilensium.
Bibliotheca Mycologica 99, Cramer, Vaduz. 339p.
Garrido N (1988) Agaricales s.l. und ihre Mykorrhizen in
den Nothofagus-Wäldern Mittelchiles. Bibliotheca
Mycologica 120: 1-528.
Grassi E, Romano G & Schenone N (2016) Macrofungi
present in a recovered zone of Atlantic Forest
(Misiones, Argentina). Boletín de la Sociedad
Argentina de Botánica 51: 223-233.
Guzman-Dávalos L (1996) New records of the genus
Gymnopilus (Agaricales, Cortinariaceae) from
Mexico. Mycotaxon 59: 61-78.
Guzmán-Dávalos L (2003) Type studies of Gymnopilus
(Agaricales) I. Mycotaxon 86: 395-423.
Guzmán-Dávalos L, Mueller GM, Cifuentes J, Miller
AN & Santerre A (2003) Traditional infrageneric
classication of Gymnopilus is not supported by
ribosomal DNA sequence data. Mycologia 95:
1204-1214.
Guzmán-Dávalos L, Contu M, Ortega A & Santerre
A (2008) New morphological and molecular data
on Gymnopilus purpureosquamulosus and its
phylogenetic relationships among similar species.
Sydowia 60: 41-56.
Guzmán-Dávalos L & Ovrebo CL (2001) Some species
of Gymnopilus from Costa Rica and Panama.
Mycologia 93: 398-404.
Hesler LR (1969) North American species of Gymnopilus.
Mycological Memoir no. 3. Hafner, New York. Pp.
1-117.
Høiland K (1998) Gymnopilus purpureosquamulosus and
G. ochraceus snov. (Agaricales, Basidiomycota) two
new species from Zimbabwe. Mycotaxon 69: 81-85.
Holec J (2005) The genus Gymnopilus (Fungi,
Agaricales) in the Czech Republic with respect
to collections from other European countries.
Acta Musei Nationalis Pragae, Series B. Historia
Naturalis 61: 1-52.
Horak E (1989) New and additional data concerning
Pyrrhoglossum and eccentric or laterally stipitate
taxa of Gymnopilus (Agaricales). Opera Botanica
100: 115-129.
Karstedt F & Stümer SL (2008) Agaricales em áreas
de oresta ombróla densa e plantações de Pinus
no estado de Santa Catarina, Brasil. Acta Botanica
Brasilica 22: 1036-1043.
Kirk PM, Cannon PF, Minter DW & Stalpers JA (2008)
Ainsworth & Bisby’s dictionary of the Fungi. CAB
International, Wallingford. Pp. 1-771.
Kühner R (1984) Some mainlines of classication in the
gill fungi. Mycologia 76: 1059-1074.
Lazo W (1984) Introducción al estudio de los hongos
superiores. Boletín Micológico 2: 27-66.
Lechner BE, Wright JE & Popoff O (2006) New taxa
and new records for Argentina of fungi from
Iguazú National Park, Misiones. Fungal Diversity
21: 131-139.
Lobato RC, Rodrigues K, Silveira ES & Vargas VS
(2010) Ocorrência de fungos do gênero Gymnopilus
Karst. em dunas na Praia do Cassino Rio Grande,
RS, Brasil. Revista Eletrônica de Biologia 3: 14-19.
Lodge DJ, Ammirati JF, O’Dell TE, Mueller GM,
Huhndorf SM, Wang CJ, Stokland JN, Schmit JP,
Ryvarden L, Leacock PR, Mata M, Umaña L, Wu Q
& Czederpiltz DL (2004) Terrestrial and lignicolous
macrofungi. In: Mueller GM, Bills GF & Foster
MS (eds.) Biodiversity of Fungi. Inventory and
monitoring methods. Elsevier Academic Press,
Cambridge. 777p.
Magnago AC, Oliveira JSS, Furtado ANM, Urrea-
Valencia S & Neves MA (2013) Mushrooms.
cogumelos. In: Neves MA, Baseia IG, Drechsler-
Santos ER & Góes-Neto A (eds.) Guide to the
common fungi of the Semiarid Region of Brazil.
TECC Editora, Florianópolis. Pp. 1-142.
Matheny PB, Curtis JM, Hofstetter V, Aime MC,
Moncalvo JM, Ge ZW, Yang ZL, Slot JC, Ammirati
JF, Baroni TJ, Bougher NL, Hughes KW, Lodge
DJ, Kerrigan RW, Sidl MT, Aanen DK, DeNitis M,
Daniele GM, Desjardin DE, Kropp BR, Norvell
LL, Parker A, Vellinga EC, Vilgalys R & Hibbett
DS (2006) Major clades of Agaricales: a multilocus
phylogenetic overview. Mycologia 98: 982-995.
Murrill WA (1913) The Agaricaceae of tropical North
America: VI. Mycologia 5: 18-36.

Gymnopilus in Paraguay 13 de 13
Rodriguésia 72: e00752019. 2021
Neves MA, Baseia IG, Drechsler-Santos ER & Góes-
Neto A (2013) Guide to the common Fungi of
the Semiarid Region of Brazil. TECC Editora,
Florianópolis. Pp. 1-142.
Niveiro N, Popoff OF & Albertó EO (2012) Presence
of Leucocoprinus cretaceus and L. fragilissimus in
Argentina. Mycotaxon 121: 265-273.
Patouillard NT & Gaillard A (1888) Champignons du
Vénézuéla et principalement de la région du Haut-
Orénoque, récoltés en 1887 par M.A. Gaillard.
Bulletin de la Société Mycologique de France 4:
7-46.
Saccardo PA (1887) Sylloge fungorum. Patavii 5: 1-1146.
Sede SM & López SE (1999) Xylophagous fungi of
urban trees in Buenos Aires City. Mycologist 13:
173-175.
Sequeira A (2013) Hongos. Guía visual de especies de
Uruguay. Ediciones de la Plaza, Montevideo. 397p.
Silva-Junior FCS & Wartchow F (2015) Gymnopilus
purpureograminicola (Strophariaceae,
Agaricomycetidae), a new species from Paraíba,
Brazil. Nova Hedwigia 101: 395-402.
Singer R (1951) Type studies on agarics III. Lilloa 25:
463-514.
Singer R (1953a) Type studies on Basidiomycetes VI.
Lilloa 26: 57-159.
Singer R (1953b) Quelques agarics nouveaux de
l’Argentine. Revue de Mycologie 18: 3-23.
Singer R (1961) Diagnoses Fungorum novorum
Agaricalium II. Sydowia 15: 45-83.
Singer R (1969) Mycoflora australis. Beih Nova
Hedwigia 29: 1-405.
Singer R (1975) Interesting and new species of
Basidiomycetes from Ecuador. Cramer, Vaduz.
Pp. 1-106.
Singer R (1986) The Agaricales in modern taxonomy.
Koeltz, Koenigstein. Pp. 1-916.
Singer R & Digilio APL (1951) Pródromo de la ora
agaricina argentina. Lilloa 25: 5-461.
Pegler DN (1983) Agaric ora of the Lesser Antilles.
Kew Bulletin Additional Series 9: 1-668.
Pegler DN (1988) Agaricales of Brazil described by M.J.
Berkeley. Kew Bulletin 43: 453-473.
Pegler DN (1990) Agaricales of Brazil described by
J.P.F.C. Montagne. Kew Bulletin 45: 161-177.
Pegler DN (1997) The agarics of São Paulo, Brazil. Royal
Botanic Gardens, Londres. Pp. 1-70.
Piris Da Motta F, Martínez M & Lechner B (2015)
Gymnopillus subtropicus Hesler (Agaricales,
Strophariaceae): nuevo género y nueva especie para
el Paraguay. Steviana 7: 84.
Putzke J (1994) Lista dos fungos Agaricales
(Hymenomycetes, Basidiomycotina) referidos
para o Brasil. Caderno de pesquisa, Série botânica
6: 1-189.
Spegazzini C (1889) Fungi Puiggariani. Pugillus 1.
Boletín de la Academia Nacional de Ciencias en
Córdoba 11: 381-622.
Spegazzini C (1919) Reliquiae mycologicae tropicae
et fungi Costaricenses nonnulli. Boletín de la
Academia Nacional de Ciencias Córdoba 23:
365-609.
Talice RV & Talice ML (1980) Hongos comestibles de
la América Meridional. Poligraf, Montevideo. Pp.
1-112.
Vasco-Palacios AM & Franco-Molano AE (2013)
Diversity of Colombian macrofungi (Ascomycota-
Basidiomycota). Mycotaxon 121: 100-158.
Wright JE & Albertó E (2002) Guía de los hongos de
la región Pampeana. I. Hongos con laminillas.
L.O.L.A., Buenos Aires. 279p.
Wright JE, Lechner BE & Popoff OF (2008) Hongos.
Atlas pictórico del Parque Nacional Iguazú.
L.O.L.A., Buenos Aires. 227p.
Wright JE & Wright E (2005) Checklist of the mycobiota
of Iguazu National Park (Misiones, Argentina).
Boletín de la Sociedad Argentina de Botánica 40:
23-44.
Area Editor: Dr. Mauricio Salazar-Yepes
Received in June 19, 2019. Accepted in November 22, 2019.
This is an open-access article distributed under the terms of the Creative Commons Attribution License.