MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 Mycotaxon, Ltd. ©2017
October–December 2017—Volume 132, pp. 819–829
https://doi.org/10.5248/132.819
Colpoma guadueticola sp. nov. in a guadual forest
from Quindio Department, Colombia
Tania Raymundo1, Ricardo Soto-Agudelo2 & Ricardo Valenzuela1*
1
Instituto Politécnico Nacional, Escuela Nacional de Ciencias Biológicas,
Plan de Ayala y Carpio s/n Col. Santo Tomas,
Ciudad de México, CP 11340, México
2
Docente del Programa de Biología, Universidad del Quindío,
Armenia, Departamento del Quindío, Colombia
* Correspondence to: rvalenzg@ipn.mx
Abstract—Colpoma guadueticola is described as a new species growing on fallen culms
of Guadua angustifolia from Colombia. The specimens were collected in a guadual forest
from the Natural Reserve ‘La Montaña del Ocaso’, municipality of Quimbaya, Quindio
Department, Colombia. A key to the species of Colpoma is presented.
Key words—Leotiomycetes, Rhytismatales, Rhytismataceae, taxonomy
Introduction
Colpoma was described by Wallroth in 1833 with C. quercinum (Pers.) Wallr.
as the type species (Darker 1967). The ascomycete genus is characterized by
its elongated (often curved) or ellipsoid apothecial ascomata that erupt under
periderm or break through naked wood and open with a single longitudinal
slit, well developed thick dark colored covering and basal layers, abundant
slender paraphyses that often coil or bend at the tip, and continuous cylindric
or filiform ascospores (Darker 1967, Johnston 1991). Quélet placed Colpoma
within Patellariaceae (Hysteriales), a family later transferred to Phacidiales by
Saccardo (Theon 1918). Darker (1967) moved Colpoma to Hypodermataceae
(Phacidiales) after which Sherwood (1979) and Johnston (1991) transferred the
genus to Rhytismataceae (Phacidiales). Currently, Medardi (2002, 2004) and
Kirk et al. (2008) include Rhytismataceae within Rhytismatales.
820 ... Raymundo, Soto-Agudelo & Valenzuela
The Natural Reserve ‘La Montaña del Ocaso’ is located at 4°34′08′′N
75°51′03′′W, south of Quimbaya municipality in Quindio Department,
Colombia (Agudelo & Gómez 2001). The guadual forest of this natural
reserve is a floristic resource of great ecological importance because of
the huge diversity of plants, animals, and fungi. The forest is dominated
by Guadua angustifolia Kunth (Poaceae), known as “Colombian bamboo”
and considered the world′s third largest bamboo, reaching heights up to 30
meters (Agudelo & Gómez 2001, Moreno et al. 2007).
There are few studies on fungal species associated with Guadua angustifolia
in Colombia. Restrepo-F. et al. (2005) isolated four agaricomycete genera
on Guadua: Pleurotus spp., Schizophyllum commune Fr., Daedaleopsis
tricolor (Bull.) Bondartsev & Singer, and Polyporus sp. Soto-Agudelo et al.
(2016a) described 29 ascomycete species collected in a guadual forest from
the Natural Reserve ‘La Montaña del Ocaso’ from Colombia; of these, 15
grew on decayed debris of Guadua angustifolia, ten were found on decayed
wood of trees, and four were parasites of other ascomycetes. In 2015 and
2016, during an ongoing survey of Ascomycetes in Colombia (Soto-Agudelo
et al. 2016a,b), three Colpoma specimens were gathered from Guadua
angustifolia. These collections had a distinct morphotype that could not
be satisfactorily accommodated in any of the known species of this genus.
The objective of the present work is to describe these specimens as a new
species of Colpoma associated with Guadua angustifolia in the Department
of Quindio in Colombia.
Materials & methods
The specimens were collected in The Natural Reserve ‘La Montaña del Ocaso’in
October 2015 and May 2016 and are deposited in the herbarium of the University
of Quindío, Armenia, Colombia (HUQ) and in the fungal collection ‘Dr. Gastón
Guzmán Huerta’ at the Herbarium Escuela Nacional de Ciencias Biológicas of Instituto
Politécnico Nacional, Mexico City, Mexico (ENCB). Latitude/longitude coordinates
were obtained with GPS eTrex (Garmin). Colors are coded according to Kornerup
& Wanscher (1978). Morphological examinations were conducted using protocols
outlined by Johnston (1991) and Minter (1997). Anatomical characters were measured
from rehydrated tissues in 5% aqueous KOH and Melzer′s reagent. Macroscopic
characters were photographed with a Nikon D7000 and micrographs were taken with a
Nikon Coolpix 9000. The meanings of some terms are based on Ulloa & Hanlin (2012)
dictionary.
Colpoma guadueticola sp. nov. (Colombia) ... 821
Figs 1–4: Colpoma guadueticola (isotype, ENCB Raymundo 5898). 1. Gregarious linear
apothecia on fallen culm. 2. Mature fusiform, curved, sigmoid, sinuous to undulate apothecia
with rounded or acute ends. 3. Elongated apothecium with rounded ends. 4. Apothecia ellipsoid,
elongated, lacrymoid with rounded or acute ends. Scale bars: 1 = 10 mm; 2 = 3 mm; 3, 4 = 1 mm.
822 ... Raymundo, Soto-Agudelo & Valenzuela
Taxonomy
Colpoma guadueticola Raymundo, Soto-Agudelo & R. Valenz., sp. nov.
Figs 1–15
MycoBank MB 820664
Differs from Colpoma quercinum by its long, filiform ascospores, its paraphyses
dichotomically to irregularly branched at the apex, and its substrate of fallen culms of
Guadua angustifolia.
Type: Colombia, Quindio department: municipality of Quimbaya, Natural Reserve ′La
Montaña del Ocaso′, 4°34′08″N 75°51′03″W, alt. 1050 m, 29 October 2015, T. Raymundo
5898 (Holotype, HUQ; isotype, ENCB).
Etymology: Referring to Guaduetum, the phytosociological name for the guadual
forest dominated by Guadua angustifolia, where the specimens were collected.
Apothecia (1–)2–10 mm long and 0.4–0.8 mm broad, stromatic, erumpent,
deeply immersed within substratum, and aligned with the main axis of the
fallen culm; visible initially as small ellipsoid to oblong or linear black spots
when closed, later opening by one longitudinal split, variable in shape when
mature: ellipsoid, oblong, fusiform, curved, sigmoid, lacrymoid, sinuous to
undulate, rounded or acute at both ends; the apothecial wall mostly covered
with black stromatic tissue. Hymenium slightly raising the substratum
surface, yellowish white (4A2) to pale yellow (4A3) in young specimens
or at the edges and ends of the ascomata, light yellow (4A5) to deep yellow
(4A8) in mature fresh specimens. Stromatic tissue in vertical transverse
section covering the lateral sides and the lower part of the apothecia,
lateral walls 60–100 µm thick, forming a textura angularis, dark brown to
black in KOH, comprising mostly angular cells, olivaceous green to black,
slightly thick-walled 5–9 µm diam.; lower wall 30–40 µm thick, forming a
textura angularis, dark brown in KOH, comprising mostly angular cells,
olivaceous green to brown, slightly thick-walled, 5–7 µm diam. Medullar
excipulum 30–50 µm thick, forming a textura intricata with interwoven
hyphae, hyaline in KOH, septate, 1–2 µm diam. Subhymenium 8–12 µm
thick, forming a textura intricata, interwoven hyphae hyaline in KOH, 1–2
µm diam. Paraphyses filiform, hyaline in KOH, non-septate, smooth, thin
walled, dichotomously to irregularly branched at the apex, extending up to
20 µm beyond the asci, 1–2 µm diam. Asci 136–180 × 5–7 µm, cylindrical,
Figs 5–11: Colpoma guadueticola (isotype, ENCB Raymundo 5898). 5. Apothecium, vertical
transverse section. 6. Hymenium. 7. Medullary excipulum, hymenium, and stromatic tissue.
8. Lateral stromatic tissue with textura angularis. 9. Basal stromatic tissue. 10–11. Basal and lateral
stromatic tissue and medullary excipulum. Scale bars: 5 = 300 µm; 6 = 80 µm; 7 = 100 µm; 8 = 40
µm; 9, 10 = 50 µm; 11 = 20 µm.
Colpoma guadueticola sp. nov. (Colombia) ... 823
824 ... Raymundo, Soto-Agudelo & Valenzuela
8-spored, hyaline in KOH, uniformly thin-walled, with only one wall layer
visible, rounded apex, inamyloid. Ascospores parallel to subparallel within
the ascus, (68–)80–110(–120) × 0.8–1.0 µm, filiform, straight to slightly
curved, hyaline in KOH, aseptate, thin-walled, smooth, gelatinous sheath
not observed at the apex.
Ecology & distribution—Gregarious on fallen and dead culms of
Guadua angustifolia in guadual forest mixed with tropical dry forest. The
species is known only from the type locality.
Additional specimens examined—COLOMBIA, Quindio department:
Municipality of Quimbaya, Natural Reserve ‘La Montaña del Ocaso’, 4°34′08″N
75°51′03″W, alt. 1050 m, 29 October 2015, R. Soto-Agudelo 60 (HUQ); 17 May 2016,
T. Raymundo 6036 (HUQ).
Comments—Colpoma guadueticola is characterized by its variably shaped
apothecia with pointed or rounded ends, yellow hymenium, paraphyses
that are dichotomously to irregularly branched at the apex, long straight to
slightly curved aseptate ascospores, and its host Guadua angustifolia.
Colpoma azaleae (Schwein.) Cooke, C. morbidum (Peck) Sacc., C. quercinum,
and C. styriacum Remler resemble C. guadueticola in having ascospores
longer than 70 µm, but C. azaleae is separated by its small (1–3 mm
long) apothecium, dark subrufescent hymenium, and Rhododendron
periclymenoides (Ericaceae) substrate; C. morbidum is distinguished by its
white to gray hymenial surface and growth on Picea mariana (Pinaceae) and
Chamaecyparis thyoides (Cupressaceae); C. quercinum has roughly circular
to oblong apothecia, apices of the paraphyses developing curls and spirals,
and growth on Quercus spp.; and C. styriacum is separated by its small
oblong yellowish green apothecia, 7–8-septate ascospores, and growth on
Vaccinium myrtilloides (Ericaceae). The other Colpoma species are separated
by their shorter (≤75 µm) ascospores and different hosts, as shown in the
key below.
Key to the species of Colpoma
1. Ascospores ellipsoid to oblong-elliptical, 5–10 µm diam., 0–3-septate . . . . . . . . . . . . 2
1. Ascospores filiform, 1–3 µm, diam., 0–8-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. On fallen bark of Agathis australis
(apothecia 0.6–1 × 0.3–0.5 mm, broad ellipsoid to oval, hymenial surface
Figs 12–15: Colpoma guadueticola (isotype, ENCB Raymundo 5898). 12. Paraphyses
dichotomously to irregularly branched at the apex. 13. Asci and paraphyses. 14. Asci with parallel
to subparallel ascospores. 15. Ascospores. Scale bars: 12, 14 = 30 µm; 13, 15 = 25 µm.
Colpoma guadueticola sp. nov. (Colombia) ... 825
826 ... Raymundo, Soto-Agudelo & Valenzuela
yellow to orange, ascospores 17–25 × 5–7.5 µm, ellipsoid, (0–)1 septate;
known only from New Zealand; Johnston 1991: 405) . . . . . . . . . . . . . . . . C. agathidis
2. On fallen wood of Nothofagus
(apothecia 1.5–5 × 1–2.5 mm, ellipsoid to irregular in shape,
hymenial surface translucent yellow, ascospores 31–46 × 6–10 µm, ellipsoid,
(0–)3-septate; known only from New Zealand; Johnston 1991: 407) . . C. nothofagi
3. Ascospores ≤75 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Ascospores >75 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
4. On conifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4. On angiosperms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
5. On Cupressaceae
(on living and dead branches of Juniperus communis, apothecia
1–3 × 1–2 mm, circular, ellipsoid to irregular in shape, hymenial surface
pale gray to fawn, ascospores 40–50 × 1–2 µm, almost straight, aseptate;
European species; Minter 1996a: 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. juniperi
5. On Pinaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Paraphyses unbranched, straight, slightly swollen at the apex
(apothecia small, 0.6–1 × 0.28–0.4 mm, long ellipsoid, round or acute at
each end, hymenial surface gray to dark gray, ascospores 33–53 × 2.5–3.5 µm,
fili-fusiform, 3-septate at maturity; known only from China on dead or
senescent twigs of Pinus armandii; Hou & Piepenbring 2005: 360) . . C. intermedium
6. Paraphyses unbranched or branched, not swollen at the apex
(circinate or flexuous with hooked tips, apothecia larger than 1 mm) . . . . . . . . . . . 7
7. Apothecia with crenulate margin
(1–2 × 0.5–0.75 mm, ellipsoid, subglobose to irregularly oblong and flexuous,
hymenial surface grayish blue to violaceus gray, ascospores 70–75 × 2 µm,
almost straight, rounded apex and basal end acute, aseptate; known only
from Sweden on fallen twigs of Pinus sylvestris; Saccardo 1891: 1128) . . C. serrulatum
7. Apothecia with smooth, striate or rugose margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Paraphyses branched, circinate
(apothecia 1–4 × 0.7–1 mm, ellipsoid to linear, hymenial surface pale
yellowish gray, ascospores 30–45 × 1.5–2 µm, aseptate; known only
from USA on decorticated conifer wood; Sherwood 1979: 35) . . . . . . . . C. deustum
8. Paraphyses unbranched, flexuous with hooked tips . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Apothecia 0.7–3 × 0.4–1 mm, sometimes curved or irregularly branched,
hymenial surface grayish to bluish gray (ascospores 26–42 × 1.5–2 µm,
rod-shaped to filiform, aseptate; European species, growing on fallen twigs
of Picea abies; Minter 1997: 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. crispum
9. Apothecia 2–3 mm long, oblong to linear, flexuous, black,
(ascospores 25–35 × 1–1.5 µm, subfiliform, rounded apex and acute base,
aseptate; known only from Sweden, growing on bark of fallen twigs
of Picea abies; Saccardo 1891: 1127) . . . . . . . . . . . . . . . . . . . . C. pseudographioides
Colpoma guadueticola sp. nov. (Colombia) ... 827
10. Apothecia with milk white hymenial surface
(ascospores 50–75 µm long, filiform; known only from North America
on dead branches of Rhododendron groenlandicum; Peck 1876: 69,
Saccardo 1883: 804) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lacteum
10. Apothecia with gray, bluish gray or grayish blue hymenial surface . . . . . . . . . . . . 11
11. Paraphyses unbranched, straight to flexuous, apex not differentiated,
apothecia small, ≤1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11. Paraphyses unbranched or branched, coiled or convoluted at the apex,
apothecia >2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12. Apothecia, 0.7–1.2 × 0.4–0.6 mm ellipsoid to almost circular
(paraphyses straight, ascospores 35–62 × 2–3 µm, fili-fusiform, tapering
towards the base, aseptate; known only from China on dead or senescent twigs
of Rhododendron sp.; Hou & Piepenbring 2005: 363) . . . . . . . . . . . . . . . . C. sinense
12. Apothecia 0.5–1.5 × 0.25–0.35 mm, oblong
(paraphyses straight to flexuous, ascospores 35–70 × 1.5–2 µm, filiform,
straight to slightly curved, aseptate; known only from China on dead branches
of Rosa sp.; Trotter 1972: 458 as Lophodermium rosae Teng) . . . . . . . . . . . . C. rosae
13. Apothecia small (0.5–5 × 0.4–1 mm)
(elliptical to oblong elliptical, aligned with the main axis of the substratum,
ascospores 33–55 × 1–2 µm, almost straight, aseptate; European species,
on dead attached twigs of Rhododendron tomentosum; Minter 1996b: 1) . . . . . . C. ledi
13. Apothecia much larger (30–35 × 2–3 mm)
(irregularly oblong, tubular, curved, sigmoid, sometimes also with bifurcations,
ascospores 60–75 × 1–1.5 µm, slightly curved, aseptate; known only from Italy
on dead attached twigs of Castanea; Medardi 2004: 36) . . . . . . . . . . . . . . C. caesium
14. On dead wood of conifers,
(apothecia ellipsoid, oblong to orbicular, white to gray in hymenial surface,
ascospores 75–100 × 1.5 µm, filiform, only known from boreal America on
Picea mariana and Chamaecyparis thyoides; Saccardo 1883: 803, Ellis & Everhart
1892: 723 as Clithris morbida (Pk.) Ellis & Everhart) . . . . . . . . . . . . . . C. morbidum
14. On angiosperms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Paraphyses dichotomously to irregularly branched at the apex
(apothecia 2–10 × 0.4–0.8 mm, ellipsoid, oblong, fusiform, curved, sigmoid,
lacrymoid, sinuous to undulate, round or acute at both ends, hymenial surface
light yellow to deep yellow, on fallen and dead culms of Guadua angustifolia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. guadueticola
15. Paraphyses unbranched, circinate or coiled at the apex . . . . . . . . . . . . . . . . . . . . . . 16
16. On Fagaceae and living/dead branches of Quercus spp.
(apothecia 2–15 × 0.5–3 mm, fusiform, navicular to curved, flexuous,
hymenial surface yellow, greenish yellow to olivaceous yellow, ascospores
70–95 × 1–3 µm, slightly curved, multiseptate, paraphyses curled to coiled,
hooked; European species; Saccardo 1883: 722, Dennis 1981: 226) . . . . C. quercinum
16. On Ericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
828 ... Raymundo, Soto-Agudelo & Valenzuela
17. On dead bark of Rhododendron periclymenoides
(apothecia 1–3 mm long, linear to elongate, flexuous, hymenial surface
dark red, ascospores 90 × 2 µm, filiform, aseptate).
Known only from boreal America (Cooke 1889: 58, Saccardo 1891: 1128,
Ellis & Everhart 1892: 722 as Clithris) . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. azaleae
17. On fallen dead twigs of Vaccinium myrtilloides
(apothecia 1–2 × 0.6–1 mm, elongate, hymenial surface translucent yellowish
green, ascospores 64–91 × 2–3 µm, sigmoid, attenuated base, 7–8 septate).
Known only from Austria (Remler 1980: 173) . . . . . . . . . . . . . . . . . . C. styriacum
Acknowledgments
We wish to express our gratitude to Dr. Peter R. Johnston and Dr. Martín Esqueda
for their useful comments and manuscript review. The authors thank the authorities
of Natural Reserve “La montaña del Ocaso” and University of Quindio for the support
and facilities to carry out the present study. Tania Raymundo thanks IPN for financial
research support in the projects SIP-20161166 and SIP-20170846. Ricardo Valenzuela
thanks COFAA and IPN for financial research support in the projects SIP-20161164
and SIP-20170845.
Literature cited
Agudelo CA, Gómez GD. 2001. Reserva Natural La Montaña del Ocaso: un nuevo modelo de
conservación. 26–41, in: Monografías de la flora andina. Importancia de la microcuenca del río
Roble. Armenia: Conceptos gráficos. Ltd. Universidad del Quindío.
Cooke MC. 1889. Memorabilia. Grevillea 17(83):58–59.
Darker GD. 1967. A revision of the genera of the Hypodermataceae. Canadian Journal of Botany 45:
1399–1444. https://doi.org/10.1139/b67-145
Dennis RWG. 1981. British Ascomycetes. J. Cramer, Vaduz. 585p.
Ellis JB, Everhart BM. 1892. The North American pyrenomycetes. USA, New Jersey, Newfield.
793 p.
Hou CL, Piepenbring M. 2005. Two new species of Colpoma on trees from China. Forest Pathology
35: 359–364. https://doi.org/10.1111/j.1439-0329.2005.00415.x
Johnston PR. 1991. Rhytismataceae in New Zealand 5. Wood- and bark-inhabiting species
in the genera Colpoma and Propolomyces. New Zealand Journal of Botany 29: 405–410.
https://doi.org/10.1080/0028825X.1991.10415493
Kirk PM, Canon PF, Minter DW, Stalpers JA. 2008. Ainsworth & Bisby′s dictionary of the fungi. 10
ed. International Mycological Institute, CAB International, Wallingford. 771 p.
Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. 3rd ed. Eyre Methuen, London.
Medardi G. 2002. Studio sul genere Colpoma Wallroth in Italia. Rivista di Micologia 45(3): 239–250.
Medardi G. 2004. Colpoma caesium: una nuova specie di Colpoma dal l′Italia. Rivista di Micologia
47(1): 35–41.
Minter DW. 1996a. Colpoma juniperi. IMI Descriptions of Fungi and Bacteria 1293. 3 p.
Minter DW. 1996b. Colpoma ledi. IMI Descriptions of Fungi and Bacteria 1294. 3 p.
Minter DW. 1997. Colpoma crispum. IMI Descriptions of Fungi and Bacteria 1333. 3 p.
Moreno LE, Trujillo EE, Osorio LR. 2007. Estudio de las características físicas de haces de fibra de
Guadua angustifolia. Scientia et Technica Año XIII, 34: 613–617.
Colpoma guadueticola sp. nov. (Colombia) ... 829
Peck CH. 1876. Report of the Botanist (1874). Annual Report on the New York State Museum of
Natural History 28: 31–88.
Remler P. 1980. Ascomyceten auf Ericaceen in den Ostalpen. Bibliotheca Mycologica 68. 321 p.
Restrepo-F GM, Vélez-A PE, Botero-A PA, Pulido-V C. 2005. Reconocimiento de macromicetos
asociados al cultivo de Guadua angustifolia en Caldas, Colombia. Manejo Integrado de Plagas
y Agroecología 76: 25–31.
Saccardo PA. 1883. Sylloge pyrenomycetum. Sylloge Fungorum 2. 813 p.
Saccardo PA. 1891. Supplementum universale. Sylloge Fungorum 9. 1141 p.
Sherwood MA. 1979. New Phacidiales and Ostropales from the collection of the Farlow herbarium.
Occasional Papers of the Farlow Herbarium of Cryptogamic Botany. 14: 35–43.
Soto-Agudelo R, Valenzuela R, Gómez-Marín GD, Raymundo T. 2016a. Ascomicetos de los
guaduales en la reserva natural La Montaña del Ocaso, departamento del Quindío, Colombia.
Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 40: 438–449.
https://doi.org/10.18257/raccefyn.352
Soto-Agudelo R, Valenzuela R, Gómez-Marín GD, Raymundo T. 2016b. Dos nuevos registros de
micromicetos saprobios del Quindío, Colombia. Boletín de la Sociedad Micológica de Madrid
40: 9–12.
Tehon LR. 1918. Systematic relationship of Clithris. Botanical Gazette 65(6): 552–555.
https://doi.org/10.1086/332288
Trotter A. 1972. Supplementum universale pars XI. Sylloge Fungorum 26. 1563 p.
Ulloa M, Hanlin RT. 2012. Illustrated dictionary of mycology. 2nd ed. APS Press, St. Paul,
Minnesota, USA. 761 p.