the journal of biodiversity data

“© Check List > PENSUFT.

ANNOTATED LIST OF SPECIES Check List 17 (5): 1383-1393

https://doi.org/10.15560/17.5.1383

New records of Agaricaceae (Agaricales, Basidiomycota) from Colombia

Jorge A. Luna-Fontalvo", Alejandra Barrios', Cristian Abaunza', Kevin Ramirez-Roncallo!, Maria A. Negritto!

1 Grupo de Conservacion y Manejo de Fauna, Flora y Ecosistemas Estratégicos Neotropicales — MIKU, Universidad del Magdalena, Santa Marta, Magdalena, Colombia + JALF: jluna@unimagdalena.edu.co @ https://orcid.org/0000-0001-6764-2276 « AB: alejandrabarriosy@unimagdalena. edu.co ® https://orcid.org/0000-0002-0938-6355 * CA: cristianabaunzafs@unimagdalena.edu.co ® https://orcid.org/0000-0002-0938-6355 * KRR: kramirezroncallo@gmail.com ® https://orcid.org/0000-0002-9462-7626 * MAN: mnegritto@unimagdalena.edu.co ® https://orcid. org/0000-0002-2508-0065

* Corresponding author

Abstract

The Agaricales are the largest clade of mushroom-forming fungi, in Colombia, with 544 species of the approximately 16,000 species currently recognized worldwide in the phylum Basidiomycota. We document seven species of Agari- caceae for the first time from the Colombian Caribbean region. Two of these species are newly recorded from the country: Leucoagaricus lilaceus Singer and Leucoagaricus roseilividus (Murrill) E. Ludw. Five species are reported from the Colombian Caribbean for the first time: Agaricus griseorimosus Pegler, Chlorophyllum hortense (Murrill) Vellinga, Leucoagaricus rubrotinctus (Peck) Singer, Leucocoprinus cepistipes (Sowerby) Pat., and L. venezuelanus Dennis.

Keywords Biodiversity, Macromycetes, macrofungi, tropical dry forest, urban reserve

Academic editor: Ajay Kumar Gautam | Received 2 July 2021 | Accepted 19 September 2021 | Published 8 October 2021

Citation: Luna-Fontalvo JA, Barrios A, Abaunza C, Ramirez-Roncallo K, Negritto MA (2021) New records of Agaricaceae (Agaricales, Basidiomycota) from Colombia. Check List 17 (5): 1383-1393. https://doi.org/10.15560/17.5.1383

Introduction

The Agaricales (Agaricomycetes, Basidiomycota) con- stitute the largest clade of mushroom-forming fungi, including 40 families, 374 genera, and more than 16,000 species (He et al. 2019; Phookamsak et al. 2019; Wijay- awardene et al. 2020). The sexual reproduction structures or basidiomata of the Agaricales are characteristically cap-, umbrella-, or shelf-shaped and with a laminar, porous, or venous hymenophore (Lodge et al. 2004). The taxonomy is determined by the morphology of the hymenophores arranged in sheets that include differential

macroscopic characteristics such as size, shape, and tex- ture of each part of the pileus; thickness, edge, color, arrangement, and insertion of the sheets at the foot; mor- phology, color, appearance, and consistency of the foot; presence or absence of a ring, volvas, or curtain; spore color, in addition to organoleptic qualities (Largent 1986; Singer 1986). Additionally, among microscopic charac- teristics, the number, morphology, ornamentation, and coloration of the basidiospores are used, as are the shape and location of cystidia, the arrangement of the hyphae

©The authors. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

1384

of pileipellis, and reactions against Melzer’s reagent, KOH, and ammonia Congo red (Pegler 1983).

In Colombia, Agaricales is the order of fungi with the greatest number of species, with 544 known (Putzke et al. 2020; Vasco-Palacios and Franco-Molano 2021) of the total of 16,071 macrofungi of the phylum Basid- iomycota worldwide (He et al. 2019; Wijayawardene et al. 2020). Most species, 83%, are recorded from the Andean region, followed by the Caribbean region with 6%; 11% is distributed in the rest of the country (Franco- Molano and Uribe-Calle 2000; Franco-Molano et al. 2010; Vasco-Palacios and Franco-Molano 2013, 2021). In the Caribbean region, 139 species have been recorded, 127 in the northwestern slope of the Sierra Nevada de Santa Marta and only 12 species in Cesar (Palacio et al. 2015; Vasco-Palacios and Franco-Molano 2013). Here, we report seven Agaricales species from the Colombian Caribbean for the first time. Two of these species are newly recorded from the country. Additionally, we pro- vide illustrations and notes on the characteristics, distri- bution, and habitat of these species.

Study Area

Our study was located on the campus of the Universidad del Magdalena (Santa Marta, Magdalena, Colombia), which has an area of 52 ha and consists of remnants of native tropical dry forest vegetation (Fig. 1). The study area 1s 3.5 ha, at 11°13'18”"N, 074°1111"W.

This part of Colombia has a bimodal rain regime with

T413'0"W 74120" 74°g'o"'w

11°18'0"N

11°15°0"N

11°42'0"N

UNIVERSIDAD DEL MAGDALENA

Legend

Manzanares river Roads

Pathways

Urban reserve of TDF Water bodies University boundaries Buildings

Sports venues

11°9'0"N

74°41 5'0"W 74°12'0"W 74°o'0"W

= b i =

Mal 0 O04 0.08

Onrapeesscrrey T4420" W yer

Check List 17 (5)

two wet seasons (May—June and September—November) and two dry seasons (December—April and June—August). The annual average rain and temperature are 578 mm and 29 °C, respectively, and the mean relative humidity is 74%, with a semi-arid climate with a high water deficit in the dry season (Rangel and Carvayal 2012).

Methods

We collected of biological material in the rainy seasons from October 2018 to October 2019, following the oppor- tunistic method (Lodge et al. 2004). The specimens were documented and preserved according to standard meth- ods (Lodge et al. 2004), and the type of habitat and asso- ciated substrate were recorded. With the fresh material, the characters of taxonomic interest such as basidiomata and spore staining were recorded. Each specimen was photographed with a Nikon D7000 digital camera, and the Methuen Handbook of Colour was used for spore staining (Kornerup and Wanscher 1978).

Microscopic analysis of the collected material was performed with a Nikon stereoscope and a Zeiss Primo Star microscope with a camera. All the measurements and colors described here were obtained from rehy- drated material with KOH 5%, alcohol 70%, and water, and Congo Red and Melzer reagent were also used. The preparations were observed with the immersion objective (100x), and the description of the structures was made using standardized techniques (Largent et al. 1977; Lar- gent and Thiers 1977; Largent 1986). Measurements of

74°6'0"'W 74° 0"W row ral

UNIVERSIDAD DEL MAGDALENA Imagen ampliada

11°18'0"N

11°18"0"N

11°12'0"N

eres oie Sas

=

—41°43'20"N

: KnreEsri-HEREeGarmin,-(¢).QpenstreetMap__ 0.32 contributors, and the GIS user community

11°9'°0"N

7] Crt ey i Ta 1ow 74°11 0-W

74°80" 74°77 0"W

Figure 1. Map of the Universidad del Magdalena campus, located in Santa Marta, Magdalena, Colombia.

Luna-Fontalvo et al. | New records of Agaricaceae from Colombia

microscopic structures were made using ToupView v. 3.7. Forty basidiospores were measured to determine the Q value (length/width ratio), the sample size was determi- nate by n = x/y (x = number of measured basidiospores, and y = number of basidiomata studied), the mean (X) of the microscopic measurements is also indicated. For the identification of the species, specialized literature was consulted (Pegler 1983; Singer 1986; Kuo 2007; Petersen and Lassoe 2012). Index Fungorum (2020) was consulted for the names of species, synonyms, and authors. The specimens were deposited at the Centro de Colecciones Bioldgicas de la Universidad del Magdalena (CBUMAG).

Results

Seven species of Agaricaceae were recorded for the first time from the Colombian Caribbean. Two species are documented here for the first time from Colombia: Leu- coagaricus lilaceus Singer and L. roseilividus (Murrill) E. Ludw. The remaining five species are newly recorded from the Colombian Caribbean region: Agaricus gris- eorimosus Pegler, Chlorophyllum hortense (Murrill) Vellinga, Leucoagaricus rubrotinnctus (Peck) Singer, Leucoprinus cepistipes (Sowerby) Pat., and L. venezu- elanus Dennis.

Agaricaceae

Agaricus griseorimosus Pegler Figure 2

Material examined. COLOMBIA — Magdalena « Santa

1385

Marta, Universidad del Magdalena; 11°13'18"N, 074° 11'09"W; 21 m alt.; 12.X.2018; J. Luna-Fontalvo et al. leg.; JL, AB & CA 005; CRBUMAG:FUN('1.

Identification. Pileus fleshy, conical to bell-shaped, flat in mature specimen, 60-70 mm in diameter, grey- ish-brown; smooth and soft surface; decurved to straight margin and rimous edge (Fig. 2A). Lamellae dark choco- late brown, free and tight, with smooth edge, and lamel- lulae (Fig. 2B). Stipe yellowish-white, 65-70 x 5-6 mm, central, cylindrical, fibrous, and hollow; annulus subapi- cal, white, double, ascending, and peronate; some speci- mens present stipes with basal torment (Fig. 2B). Trama pale yellow and dark brown, compact, and soft. Basidiospores dark brown, 4.0—5.4 x 2.6-3.8 um (X = 11.2 + 0.3984 x 3.2 + 0.3095; Q = 1.46 + 0.1596; n = 50/2), ellipsoid and thick-walled, germ pore absents (Fig. 2C). Basidia were not observed. Cheilocystidia 15—18.2 x 7-9 um, hyaline and claviform (Fig. 2D). Pileipellis composed of hyphae 2-14 um in diameter, hyaline, glo- bose, with thin walls (Fig. 2E). Taxonomic remarks. The macroscopic and microscopic features of the examined material match that described by Pegler (1983) but differ slightly 1n the size of the ba- sidiospores: 5.3—7.5 x 2.7-3.5 (5.6 + 0.4 x 3 £ 0,18) um. In A. griseorimosus, the coloration of the stipe, the size of basidiospores, and the presence or absence of chei- locystidia is variable, as in A. endoxanthus, A. moel- leri, A. moelleroides, A. punjabensis, A. tephrolepidus, and A. volvatulus. According to Parra et al. (2018), A. griseorimosus belongs to the section Xanthodermatei;

Figure 2. Basidiomata and microscopic structures of Agaricus griseorimosus (JL, AB & CA 005). A. Whole basidiomata. B. Details of the hymenium and stipe. C. Spores. D. Cheilocystidia. E. Pileipelis hyphae. Scale bars: A, B= 2 cm; C= 10 um; D, E= 20 um.

1386

Check List 17 (5)

Figure 3. Basidiomata and microscopic structures of Chlorophyllum hortense (JL, AB & CA 040). A. Apical view of the pileus. B. Details of the hymenium and stipe. C. Spores. D. Basidia. E. Cheilocystidia. Scale bars: A, B= 2 cm; C = 10 um; D, E= 20 um.

however, it presents a bordered pileus, a single annulus, and longer and narrower spores which distinguishes it from that section.

Habitat and distribution. A terrestrial species, on Iit- ter in a tropical dry forest fragment. Collected during the rainy season. Agaricus griseorimosus has been recorded in India (Farook et al. 2013), the Lesser Antilles (Ange- lini et al. 2018), and Colombia, where it was previously known from the Caqueta department (Sanchez 2003). Our record is the first from the Colombian Caribbean.

Chlorophyllum hortense (Murrill) Vellinga Figure 3

Material examined. COLOMBIA — Magdalena « Santa Marta, Universidad del Magdalena; 11°13'31"N, 074° 11'03”W; 21 m alt.; 1.X.2019; J. Luna-Fontalvo et al. leg.: JL, AB & CA 040; CBUMAG:FUN:2.

Identification. Pileus fleshy, convex to flat and slightly umbonate; 50-70 mm in diameter, white with oxide brown scales, dry surface, straight to decurved margin, and torn edge (Fig. 3A). Lamellae white, free, tight, with smooth edge and lamellulae (Fig. 3B). Stipe white with light orange spots, 70—90 x 4—6 mm, central, hollow, cy- lindrical, fibrous; annulus subapical, concolorous with the stipe, peronate, and ascending; slightly widened base with mycelial laces (Fig. 3B). Trama white, thick, abun- dant, and consistent.

Basidiospores greyish-green, 7.1-9.2 x 5.2—7.3 um, broadly ellipsoidal, germ pore absents (Fig. 3C). Basidia hyaline, 29.6-35.7 x 12.0-14.1 um (X = 8.3 + 0.4019 x 6.7 + 0.4067; Q = 1.23 + 0.057; n = 40/5), claviform, tet- rasporic and granular contents (Fig. 3D). Pleurocystidia absent. Cheilocystidia with greenish yellow vacuolar pigments, spiked, 32.2—58.6 x 9.3-16 um (Fig. 3E).

Taxonomic remarks. Chlorophyllum hortense 1s char- acterized by a small to medium-size basidiome, white with brown central scales, single annulus, spores with- out germ pore and basidia with two or occasionally four sterigmata (Vellinga 2003; Ge and Yang 2006). How- ever, Pegler (1983) described spores of C. hortense with a germ pore. Chlorophyllum hortense has similarities with C. africanum in the size of the basidiocarp and the shape of the spores and cheilocystidia. However, C. afri- canum has smaller spores and basidia with four sterig- mata (Ge et al. 2018). Our Colombian material matches the features described by Vellinga (2003), Ge and Yang (2006), and Nascimento and Alves (2014), who report spores without a germ pore.

Habitat and distribution. A terrestrial species, on scat- tered pastures. Collected in the rainy season. Chloro- phyllum hortense has a wide distribution in tropical and subtropical climates and has been recorded in the Lesser Antilles, United States (Pegler 1983), Brazil (Nasci- mento and Alves 2014; Alves et al. 2016), and Colombia (Franco-Molano et al. 2010).

Leucoagaricus lilaceus Singer Figure 4

Material examined. COLOMBIA — Magdalena « Santa Marta, Universidad del Magdalena; 11°13'18"N, 074° 11'05"W; 20 m alt.; 13.X.2019; J. Luna-Fontalvo et al. leg.; JL, AB & CA 002; CRBUMAG:FUN:3.

Identification. Pileus fleshy, convex to flat, 5|0—70 mm in diameter, white, with a wide greyish ruby scale to dark ruby, scattered throughout the cap; dry surface, straight margin, rimous to lobed edge (Fig. 4A, B). Lamellae yel- lowish-white, free, tight, ventricose, with smooth edge and lamellulae (Fig. 4C). Stipe white at the apex and

Luna-Fontalvo et al. | New records of Agaricaceae from Colombia

1387

Figure 4. Basidiomas and microscopic structures of Leucoagaricus lilaceus (JL, AB & CA 002). A. Apical view of the pileus. B. Lateral view of the basidiocarp. C. Details of the hymenium and stipe. D. Spores. E. Basidia. F. Pileipelis hyphae. Scale bars: A= 2 cm; B=4cm; C=2 cm; DiSst0: anise; F=20)nm,

light brown at the base, 50-60 x 6—8 mm, central; an- nulus fleeting and absent in mature basidiomata; base thickened, with a prominent and rounded bulb (Fig. 4C). Trama white and consistent.

Basidiospores hyaline, 5.32—7.1 <x 3.2-4.8 um (X =

6.2 + 0.4739 x 4 + 0.3703; Q = 1.51 + 0.37; n = 40/3), ellipsoid and thick-walled; germ pore absents (Fig. 4D). Basidia 20.3—30 x 6.9-11.61 um, hyaline, claviform, tet- rasporic (Fig. 4E). Pleurocystidia absent; cheilocystidia not observed. Pileipellis composed of prostrate hyphae, 4—21um wide (Fig. 4F). Taxonomic remarks. Leucoagaricus lilaceus belongs to the section Piloselli (Kthn.) Sing. (1972). The mor- phological descriptions presented correspond to those of Singer (1986), except for the absence of hyaline , fu- siform, ventricose, langeriform, and thin-walled chei- locystidia (Singer 1986; Rother and Da Silveira 2009a; Vellinga et al. 2010; Ferreira and Cortez 2012).

Leucoagaricus lilaceus 1s considered close to Lepiota roseolivida Murrill, as these species both have the pileus with violet tones, but they differ in the size and color of the stipe, which is yellowish in L. roseolivida. Also, L. lilaceus is related to Lepiota decorata Zeller but is dis- tinguished by some microscopic structures, such as cys- tidia, and L. decorata is distributed in temperate zones where it is associated with conifers (de la Fuente et al. 2018). Leucoagaricus subpurpureolilacinus ZW. Ge & Zhu L. Yang also has the pileus with violet color, but the basidiospores are larger, approximately 11 um, and the context does not change color when it is cut (Ge et al. 2015).

Habitat and distribution. A terrestrial species, solitary, sometimes scattered in tropical dry forest fragments. Collected in the rainy season. Leucoagricus lilaceus 1s newly recorded from Colombia. It was first described

in Tucuman, Argentina (Singer and Digilio 1951) and is known from Brazil (Rother and Da Silveira 2009a; Fer- reira and Cortez 2012), Paraguay (Flecha-Rivas et al. 2013), and Mexico (de la Fuente et al. 2018).

Leucoagaricus roseilividus (Murrill) E. Ludw. Figure 5

Material examined. COLOMBIA — Magdalena ¢ Santa Marta, Universidad del Magdalena; 11°13'29"N, 074° 11'14”W; 20 m alt.; 11.X.2019; J. Luna-Fontalvo et al. leg.: JL, AB & CA 050; CBUMAG:FUN:4.

Identification. Pileus convex to plane-convex, 42.6—63.9 mm in diameter, white with scales over the entire grey magenta surface, darker towards the center; decurved margin, and entire edge (Fig. 5A, B). Lamellae orangish- white, free, tight, with smooth edge and lamellulae (Fig. 5C). Stipe concolorous, darker towards the base, 53.2— 56 x 4-7 mm, central, spiked and fibrillous; annulus su- perior, white above, and grey magenta below (Fig. 5C). Trama white and consistent.

Basidiospores hyaline, 5—7.3 x 3.2-5 um (X = 5.9 + 0.4716 x 3.8 0.2559; Q = 1.55 0.1550; n = 40/1), ellip- soid, with thick wall and germ pore (Fig. 5D). Basidia 21.8—29 x 7.3—9 um, hyaline, claviform, tetrasporic, with smooth and thin walls, some exhibit granular contents (Fig. SE). Pleurocystides not observed. Cheilocystides 29.8—45.4 x 9.1-16.8, polymorphic, claviform, utriform, cylindrical, wavy, oblong, slightly ventricose, with thick- ened walls, turning hyaline to yellowish in KOH, with olive-brown vacuolar pigments (Fig. 5F). Pileipellis composed of periclinal hyphae, cylindrical, interlaced, ascending, olive-brown, with terminal elements, 24.7— 40.7 x 4.6-10.5 um (Fig. 5G).

Taxonomic remarks. The morphological features of L. roseilividus described here correspond to the original

1388

Check List 17 (5)

Figure 5. Basidiomata and microscopic structures of Leucoagaricus roseilividus (JL, AB & CA 050). A. Basidiocarp. B. Apical view of the pileus. C. Details of the hymenium and stipe. D. Spores. E. Basidia. F. Cheilocystidia. G. Pileipelis hyphae. Scale bars: A= 3 cm; B, C=2 cm; D=10 um; E, G= 20 um.

description by Murrill (1912) and the descriptions by Vel- linga (2006) and Cabrera (2015); this species has a reac- tion in ammonia, but this test was not performed in the sample examined. Leucoagaricus roseilividus exhibits delicate, small basidiomata, but the size is variable and, in some habitats, may be more robust ( Vellinga 2006).

It has been confused on countless occasions due to its close resemblance to Leucoagaricus ionidicolor Bellu & Lanzoni, but it differs in the intensity of the colors, with more bluish tones, a considerably wider pileus, and an unchanging color of the context and lamellae (Vel- linga 2006). Our sample agrees with the description by Cabrera (2015) of a Brazilian specimen, specifically the coloration of the cheilocystidia and pileipellis hyphae, which are brown in water and yellowish to hyaline in KOH, with intracellular pigments, and hyphae with fibu- lae absent.

Habitat and distribution. A terrestrial, solitary species, on grass. Collected in the rainy season. Leucoagaricus roseilividus 1s recorded from Colombia for the first time. It occurs in California, United States (Murril 1912; Vel- linga 2006), and Brazil (Cabrera 2015). Ours is the sec- ond record from South America.

Leucoagaricus rubrotinctus (Peck) Singer Figure 6

Material examined. COLOMBIA — Magdalena ¢ Santa Marta, Universidad del Magdalena; 11°13'18"N, 074°11’ 10"W; 20 m alt.; 12.VI.2019; J. Luna-Fontalvo et al. leg.: JL, AB & CA 014, CRUMAG:FUN‘5.

Identification. Pileus flat to concave; 15—20 mm in di- ameter; white, covered by tiny, red, radially arranged scales,, darker in the center; surface dry and rough; with an entire edge and straight margin (Fig. 6A). Lamellae

white, free, thick, tight, with lamellulae. Stipe 118-20 x 2—4 mm, white, cylindrical, smooth, central, fibrous; an- nulus central, ascending, and peronate; base with small red scales (Fig. 6B). Trama white and consistent.

Basidiospores hyaline, 5.3-9.2 x 3-5.7 um (X = 7.3 + 0.7235 x 4.4 + 0.6705; Q = 1.68 + 0.229; n = 40/3), ellipsoid and thick-walled; germ pore absents (Fig. 6C). Basidia 17.7—28.2 x 6.7—10 um, hyaline, claviform and tet- rasporic (Fig. 6D). Pleurocystides not observed. Cheilo- cystides 19.2—35.9 x 7.1-10.9 um, hyaline, polymorphic, fusiform-langeriform, ventricose, subcapitated (Fig. 6E). Pileipellis composed of periclinal hyphae to irregularly ascending hyphae, 7—-15.2 um wide, occasionally septate, with olive-brown vacuolar pigments (Fig. 6F).

Taxonomic remarks. The taxonomic features of our material studied correspond with the descriptions of L. rubrotinctus by Ferreira and Cortez (2012), includ- ing a reddish pileus which varies from orange to brown tones depending on the age of the basidiocarp, but re- maining always darker in the center. In immature spec- imens, the pileus displays a uniform color which, when maturing, cracks and gives the impression of small radial scales on a pale background (Ferreira and Cortez 2012). Among the microscopic features, this species presents basidiospores with an inconspicuous to absent germ pore (Verma et al. 2018).

Lepiota rubrotinctoides Murrill and Leucoagari- cus glabridiscus (Sundb.) have been related with L. rubrotinctus,; however, they present macroscopic and microscopic differences that have solved the confusion (Ferreira and Cortez 2012). Lepiota rubrotinctoides presents smaller basidiospores (7 x 3.5 um), absence of hairy scales, and more robust basidiomata (Murrill 1912), whereas L. glabridiscus 1s a smaller and fragile fungus,

Luna-Fontalvo et al. | New records of Agaricaceae from Colombia

1389

Oni

Figure 6. Basidiomata and microscopic structures of Leucoagaricus rubrotinctus (JL, AB & CA 014). A. Lateral view of the pileus. B. Details of the hymenium and stipe. C. Spores. D. Basidia. E. Cheilocystidia. F. Pileipelis hyphae. Scale bars: A=1 cm; B= 2 cm; C= 10 um; D-F = 20 um.

whose pileipellis is formed by a double layer of inter- laced hyphae (Kumar and Manimohan 2009).

Habitat and distribution. A terrestrial species, solitary, on litter, in a tropical dry forest fragment. Collected dur- ing the dry season. Our record 1s the first from the Co- lombian Caribbean region. Elsewhere in Colombia this species has been found in the departments of Antioquia and Quindio, between 1800 and 3250 m altitude (Franco- Molano et al. 2010). It is also reported from Bolivia (Mel- garejo-Estrada et al. 2020), Brazil (Ferreira and Cortez 2012), and Mexico (Terriquez-Villanueva et al. 2017).

Leucocoprinus cepistipes (Sowerby) Pat. [‘cepaestipes’ |

Figure 7

Material examined. COLOMBIA — Magdalena « Santa Marta, Universidad del Magdalena; 11°13'18"N, 074°11' 10”"W; 20 m alt.; 12.VI.2019; J. Luna-Fontalvo et al. leg.; JL, AB & CA 013; CBUMAG:FUN‘6.

Identification. Pileus convex to flat, slightly depressed in mature specimens; 20-30 mm in diameter; white, with around scale in the center in some specimens, maize-yel- low; surface floccose and striated towards the margin,

entire edge, straight margin to raised (Fig. 7A). Lamellae white, free, tight, with smooth edge and lamellae (Fig. 7B). Stipe concolorous with hymenium, 40-50 x 3-4 mm, floccose; annulus fleeting, intermediate, membra- nous, fibular, ascending, with crenate border (Fig. 7C-E). Trama white, soft, and sticky.

Basidiospores hyaline, 5.7—7. x 3.6-4.5 um (X = 6.3 + 0.3642 x 4 + 0.2477; Q = 1.54 + 0.094; n = 40/4), ellipsoid, smooth, with thick wall and germ pore (Fig. 7F). Basidia 27.7-36.4 x 7.8—9.4 um, hyaline, claviform, tetrasporic, with thin wall, lacking a basal clamp (Fig. 7G). Pleurocystidia not observed. Cheilocystidia 52—78 x 12-15 um, hyaline, with smooth and thin wall, poly- morphic, spiked, ventricose, fusiform, langeriform, and some rostrate (Fig. 7H).

Taxonomic remarks. Leucocoprinus cepistipes 1s char- acterized by a white basidiocarp covered by cottony scales, which give it a floccose appearance, and a pileus with a plicate-striated margin. Microscopically it pres- ents ellipsoid, oblong basidiospores, with a conspicuous germ pore (Vellinga 2001). Our material agrees with that presented by Rother and Silveira (2009b), in having ba- sidiospores 8.5—10 x 6-7 um, Q = 1-1.8, basidia 19-24

1390

Check List 17 (5)

Figure 7. Basidiomata and microscopic structures of Leucocoprinus cepistipes (JL, AB & CA 013). A. Lateral view of the pileus. B. Details of the hymenium. C. Stipe details. D, E. Annulus details. F. Spores. G. Basidia. H. Cheilocystidia. Scale bars: A, B= 2 cm; F = 10 um; G, H = 20 um.

x 9-10 um, pleurocystidia absent, and cheilocystidia 52— 78 x 12-15, differentiated by slightly smaller spores and slightly larger basidia.

Habitat and distribution. A terrestrial species, scat- tered, on litter, in a tropical dry forest fragment. Col- lected in the dry season. Our record is the first from the Colombian Caribbean. This species was previously re- corded from Antioquia and Meta departments at 750 m alt. (Franco -Molano et al. 2010). Leucocoprinus cepis- tipes has a pantropical distribution, and in America, it has been recorded in Mexico (Guzman 1983; de la Fuente et al. 2020), the Lesser Antilles, Martinique, Dominica, Antigua, Trinidad, Jamaica, and the Bahamas (Pegler 1983), Argentina (Campi et al. 2015), Brazil (Rother and da Silveira 2009b), and Paraguay (Campi et al. 2015).

Leucocoprinus venezuelanus Dennis Figure 8

Material examined. COLOMBIA — Magdalena ¢ Santa Marta, Universidad del Magdalena; 11°13'31"N, 074°11' 04"W; 21 m alt.; 1.X.2019; J. Luna-Fontalvo et al. leg.; JL, AB & CA 041; CBUMAG:FUN:7.

Identification. Pileus convex to flat, 40-50 mm in di- ameter, white with cognac-colored scales, fragmented radially towards the edges; striated translucent surface, with straight margin, raised in mature specimens and torn edge (Fig. 8A, B). Lamellae white, free, tight, with a slightly crenate edge and lamellulae (Fig. 8C). Stipe con- colorous with hymenium, 52—60 <x 5—6 mm, central, hol- low and clavate; annulus intermediate, white, and with tiny scales cognac-colored adhered to the margin (Fig. 8C). Trama white and thin.

Basidiospores hyaline, 6.2— 8.4 x 3.7—5.3 um (X= 7.2 + 0.50 x 4.7 + 0.37; Q = 1.53 + 0.16; n = 40/2), ellipsoids

and smooth; germ pore absent (Fig. 8D). Basidia 19.07— 25.23 x 7.9-10.72 um, hyaline, claviform, tetrasporic, and with granular contents (Fig. 8E). Pleurocystidia not observed. Cheilocystidia 26.—32.9 x 10-15 um, hyaline, fusiform to claviform (Fig. 8F).

Taxonomic remarks. Leucocoprinus venezuelanus was originally described by Dennis (1961). Our material matches with the descriptions of this species by Pegler (1983), but with a slight difference in the size of the chei- locystidia. Leucocoprinus venezuelanus usually resem- bles L. brebissonni in basidiome morphology, but the former species 1s differentiated by the presence of brown scales on the annulus margin and by the size of the ba- sidiospores (Plegler 1983; Kumar and Manimohan 2009). Leucocoprinus submontagnei also resembles L. venezu- elanus, but it differs in having brown scales and spores that are slightly smaller, no germ pores, and vermiform cheilocystidia (Deepa et al. 2006).

Habitat and distribution. A terrestrial species, scat- tered and gregarious, on litter. Collected during the rainy season. This species has a wide distribution, and it has been recorded in Caqueta department, Colombia (Sanchez 2003), the Lesser Antilles (Plegler 1983), Bra- zil (Karstedt and Sttrmer 2008), and Venezuela (Den- nis 1961).

Discussion

Seven species of Agaricaceae are newly recorded from the Colombian Caribbean, which brings the number of species to 39 for this region and 551 for Colombia. Two species, Leucoagaricus lilaceus, and L. roseilivi- dus, are reported from Colombia for the first time. The area studied in Santa Marta city is extremely small, and

Luna-Fontalvo et al. | New records of Agaricaceae from Colombia

1393)

Figure 8. Basidiomata and microscopic structures of Leucocoprinus venezuelanus (JL, AB & CA 041). A. Scales of the pileus. B. Apical view of the pileus. C. Details of the hymenium and stipe. D. Spores. E. Basidia. F. Cheilocystidia. Scale bars: A-C = 2 cm; D =10 um; E, F = 20 um.

there are still large unexplored areas. In the last 40 years, macro-fungi studies have been concentrated mainly in the Andean region, with 83% of the species from that region and only 6% from the Caribbean region (Pulido 1983; Pineda et al. 1988; Franco-Molano and Uribe-Calle 2000; Franco-Molano et al. 2010; Vasco-Palacios et al. 2013; Palacio et al. 2015; Lombana et al. 2016; Putzke et al. 2020). Therefore, we provide an important contribu- tion to the inventory of Agaricales of the Caribbean Plain biogeographic region where the fungi diversity is high and almost unexplored.

The tropical dry forest exhibits water stress, high luminosity, and high temperatures during some months of the year, in addition to characteristic acidic and nutri- ent-poor soils. These forests do not have very suitable conditions for many macro-fungi. However, fallen leaves create abundant leaf litter which forms, together with components of the soil, a rich organic substrate and provides a microclimate with appropriate conditions of humidity and temperature that favor the fruiting of fungi (Dix and Webster 1995; Palacio et al. 2015). These con- ditions are present in the permanent parcel of dry forest at the Universidad del Magdalena (Barranco Pérez et al. 2019).

Mycological studies in dry forests of the tropics are very few, and the sparse data available do not allow for determining with certainty the diversity of fungi in these forests (Palacio et al. 2015). Although our contri- bution adds important new data on the high biodiversity of macro-fungi in these ecosystems, there is too little information available to make meaningful comparisons. Inventories and long-term monitoring, supplemented with molecular tools, are necessary to expand our knowl- edge of fungi in tropical dry forests.

According to Rangel-Ch. (2015), of the extra-Andean

regions, the Caribbean region is the most biodiverse, but this estimate considered flowering plants, ferns and bryophytes, amphibians, reptiles, mammals, and lichens, but not macro-fungi. This was probably due to a lack of exploration or expert taxonomists in the field. Therefore, more study of macro-fungi in the Caribbean region is needed to better determine the biodiversity of Colombia, evaluate the importance of macro-fungi in ecosystem services, and work towards their conservation.

Acknowledgements

We express gratitude to the Research Fund FONCIEN- CIAS of Universidad del Magdalena for the financial support during the research, the Grupo de Investigacion Manejo y Conservacion de Fauna, Flora y Ecosistemas Estratégicos Neotropicales, and the anonymous review- ers of the manuscript.

This is the scientific contribution no. 21 from the Centro de Colecciones Biolégicas de la Universidad del Magdalena.

References

Alves RP, Menezes GCA, Oliveira ED, Victoria FC, Pereira AB, Albu- querque MP (2016) Chlorophyllum Massee e Macrolepiota Singer (Agaricaceae) em area do bioma Pampa, Rio Grande do Sul, Bra- sil. Neotropical Biology and Conservation 11 (3): 141-152. https:// doi.org/10.4013/nbc.2016.113.04

Angelini C, Ortiz-Santana B, Mata G, Billette C, Rojo C, Chen J, Cal- lac P (2018) The genus Agaricus in the Caribbean. Nine new taxa mostly based on collections from the Dominican Republic. Phyto- taxa 345 (3): 219-271. https://doi.org/10.11646/phytotaxa.345.3.2

Barranco Pérez W, Yepes Rapelo D, Garcia Quifiones H (2019) Espe- cies comunes del bosque seco en el campus de la Universidad del Magdalena. Universidad del Magdalena, Santa Marta, Colombia, 56 pp.

1392

Cabrera CH (2015) Leucoagaricus (Agaricaceae) na regiao sul do Brasil. MSc dissertation, Universidade Federal de Santa Catarina, Santa Catarina, Brazil, 84 pp.

Campi MG, De Madrignac BR, Flecha Rivas AMI, Niveiro N (2015) El género Leucocoprinus Pat. (Agaricaceae-Agaricommycetes) en el norte de Argentina y Paraguay. Iheringia Serie Botanica 70 (2): 309-320.

de la Fuente JI, Garcia-Jiménez J, Lopez CY, Oros-Ortega I, Vela- Hernandez RY, Guevara-Guerrero G, Garza-Ocafias F, Chay-Ca- sanova J, Ibarra-Garibay L, Bandala VM (2020) An annotated checklist of the macrofungi (Ascomycota, Basidiomycota, and Glomeromycota) from Quintana Roo, Mexico. Check List 16 (3): 627-648. https://doi.org/10.15560/16.3.627

de la Fuente JI, Hernandez-Del Valle JF, Aguirre-Acosta CE, Garcia- Jiménez J (2018) First record of Leucoagaricus lilaceus (Agarica- les: Agaricomycetes) from Mexico. Studies in Fungi 3 (1): 187— 191. https://do1.org/10.5943/sif/3/1/19

Deepa S, Vrinda KB, Pradeep CK (2006) More leucocoprinoid fungi from the Western Ghats. Mushroom Research 15 (2): 103-110.

Dennis RWG (1961) Fungi venezuelani: IV. Agaricales. Kew Bulletin (15): 67-156. https://doi.org/10.2307/4115784

Dix JN, Webster J (1995) Fungal ecology. Springer, London, United Kingdom, 549 pp. https://doi.org/10.1007/978-94-011-0693-1

Farook VA, Khan SS, Manimohan P (2013) A checklist of agarics (gilled mushrooms) of Kerala State, India. Mycosphere 4 (1): 97— 131. https://doi.org/10.5943/mycosphere/4/1/6

Ferreira AJ, Cortez VG (2012) Lepiotoid Agaricaceae (Basidiomy- cota) from S40 Camilo State Park, Parana State, Brazil. Myco- sphere 3 (6): 962—976. https://doi.org/10.5943/mycosphere/3/6/11

Flecha-Rivas AM, De Madrignac B, Campi M, Ortellado (2013) Nuevo registro de Lewcoagaricus lilaceus Singer (Agaricomyce- tes — Agaricaceae) para Paraguay. Reportes Cientificos de la FA- CEN 4: 11-14.

Franco-Molano AE, Corrales A, Vasco-Palacios A (2010) Macrohon- gos de Colombia II. Listado de especies de los 6rdenes Agaricales, Boletales, Cantharellales y Russulales (Agaricomycetes, Basidio- mycota). Actualidades Biologicas 32 (92): 89-114.

Franco-Molano AE, Uribe-Calle E (2000) Hongos Agaricales y Bole- tales de Colombia. Biota Colombiana 1| (1): 25—43.

Ge Z, Jacobs A, Vellinga E, Sysouphanthong P, Van der Walt R, Lavorato C, An YF, Yang Z (2018) A multi-gene phylogeny of Chlorophyllum (Agaricaceae, Basidiomycota): new species, new combination and infrageneric classification. MycoKeys (32): 65— 90. https://doi10.3897/mycokeys.32.23831

Ge ZW, Yang ZL (2006) The genus Chlorophyllum (Basidiomycetes) in China. Mycotaxon (96): 181-192.

Ge ZW, Yang ZL, Qasim T, Nawaz R, Khalid AN, Vellinga EC (2015) Four new species in Leucoagaricus (Agaricaceae, Basidiomycota) from Asia. Mycologia (107): 1033-1044. https://doi.org/10.3852/ 14-351

Guzman G (1983) Los hongos de la peninsula de Yucatan II. Nuevas exploraciones y adiciones micoldgicas. Biotica 8: 71—87.

He MQ, Zhao RL, Hyde KD, Begerow D, Kemler M, Yurkov A, Mck- enzie EHC, Raspé O, Kakishima M, Sanchez-Ramirez S, Vel- linga EC, Halling R, Papp V, Zmitrovich IV, Buyck B, Ertz D, Wiayawardene NN, Cui BK, Schoutteten N, Liu XZ, Li TH, Yao YJ, Zhu XY, Liu AQ, Li GJ, Zhang MZ, Ling ZL, Cao B, An- tonin V, Boekhout T, da Silva BDB, Crop ED, Decock C, Dima B, Dutta AK, Fell JW, Geml J, Ghobad-Nejhad M, Giachini AJ, Gibertoni TB, Gorjon SP, Haelewaters D, He SH, Hodkinson BP, Horak E, Hoshino T, Justo A, Lim YW, Menolli JN, MeSic¢ A, Moncalvo JM, Mueller GM, Nagy LG, Nilsson RH, Noordeloos M, Nuytinck J, Orihara T, Ratchadawan C, Rajchenberg M, Silva- Filho AGS, Sulzbacher MA, Tkaléec Z, Valenzuela R, Verbeken A, Vizzini A, Wartchow F, Wei TZ, WeiB M, Zhao CL, Kirk PM (2019) Notes, outline and divergence times of Basidiomycota. Fungal Diversity 99 (1): 105-367. https://doi.org/10.1007/s13225- 019-00435-4

Check List 17 (5)

Index Fungorum (2020) Authors of fungal names. http://www.index fungorum.org/Names/AuthorsOfFungalNames.asp. Accessed on: 2020-7-13.

Karstedt F, Stirmer S (2008) Agaricales em areas de Floresta Om- brofila Densa e plantagdes de Pinus no Estado de Santa Catarina, Brasil. Acta Botanica Brasilica 22 (4): 1036-1043. https://doi.org/ 10.1590/S0102-33062008000400014

Kornerup A, Wanscher JH (1978) Methuen handbook of colour (3 edi- tion). Eyre Methuen, London, United Kingdom, 252 pp.

Kumar TKA, Manimohan P (2009) The genera Leucoagaricus and Leucocoprinus (Agaricales, Basidiomycota) in Kerala State, In- dia. Mycotaxon 108: 385-428.

Kuo M (2007) The gilled mushrooms (“Agaricales”). http://www. mushroomexpert.com/agaricales.html. Accessed on: 2020-02-15.

Largent DL (1986) How to identify mushrooms to genus I: macro- scopic features. Mad River Press, Eureka, USA, 86 pp.

Largent DL, Johnson D, Watling R (1977) How to identify mush- rooms to genus III: microscopic features. Mad River Press, Eu- reka, USA, 148 pp.

Largent DL, Thiers HD (1977) How to identify mushrooms to genus IT: field identification of genera. Mad River Press, Eureka, USA, 32 pp.

Lodge DJ, Ammirati JF, O’Dell T, Mueller GM, Huhndorf SM, Wang CJ, Stokland J, Schmit J, Ryvarden L, Leacock P, Mata M, Umafia L, Wu QF, Czederpiltz D (2004) Terrestrial and lingnicolous mac- rofungi. In: Mueller GM, Bills GF, Foster MS (Eds.) Biodiversity of Fungi: inventory and monitoring methods. Elservier Academic Press, London, United Kingdom, 127-158. https://doi.org/10.1016/ B978-012509551-8/50011-8

Lombana P, Monterroza JA, Chamorro LF, Franco-Molano AE, Payares IR (2016) Nuevos registros de macromicetos para Colom- bia. Actualidades Bioldgicas 38 (105): 181-189. https://doi.org/10. 17533/udea.acbi.v38n105a05

Melgarejo-Estrada E, Rocabado D, Suarez ME, Maillard O, Lechner BE (2020) Checklist of Bolivian Agaricales. 2: species with white or pale spore prints. Mycotaxon 135 (1): 1-29.

Murrill WA (1911) The Agaricaceae of tropical North America. IV. Mycologia 3 (6): 271-282. https://doi.org/10.2307/3753496

Murrill WA (1912) The Agaricaceae of the Pacific Coast II. Mycologia 4 (5): 231-262. https://doi.org/10.1080/00275514.1912.12017913

Nascimento CC, Alves MH (2014) New records of Agaricaceae (Ba- sidiomycota, Agaricales) from Araripe Nacional Forest, Ceara state, Brazil. Mycosphere 5 (2): 319-332. https://doi.org/10.5943/ mycosphere/5/2/6

Palacio M., Gutiérrez Y, Franco-Molano A, Callejas-Posada R (2015) Nuevos registros de macrohongos (Basidiomycota) para Colom- bia procedentes de un bosque seco tropical. Actualidades Biologi- cas 37 (102): 319-339.

Parra LA, Angelini C, Ortiz-Santana, B, Mata G, Billete C, Rojo C, Chen J, Callac P (2018) The genus Agaricus in the Caribbean. Nine new taxa mostly based on collections from the Dominican Republic. Phytotaxa 345 (3): 219-271. https://doi.org/10.11646/ phytotaxa.345.3.2

Pegler DN (1983) Agaric flora of the Lesser Antilles. Kew Bulletin Ad- ditional Series 9: 1-668. https://doi.org/10.2307/3793124

Petersen JH, Lassoe T (2012) MycoKey. http://www.mycokey.com/, Accessed on: 2020-06-20.

Phookamsak R, Hyde, KD, Jeewon R, Bhat DJ, Jones EBG, Maha- rachchikumbura SSN, Raspé O, Karunarathna SC, Wanasinghe, DN, Hongsanan S, Doilom M, Tennakoon DS, Machado AR, Fir- mino AL, Ghosh A, Karunarathna A, Mesi¢ A, Dutta AK, Thong- bai B, Devadatha B, Norphanphoun C, Senwanna C, Wei D, Pem D, Ackah FK, Wang GN, Jiang HB, Madrid H, Lee HB, Goonas- ekara ID, Manawasinghe IS, KuSan I, Cano J, Gené J, Li J, Das K, Acharya K, Raj KNA, Lath KPD, Chethana KWT, He MQ, Duefias M, Jadan M, Martin MP, Samarakoon MC, Dayarathne MC, Raza M, Park MS, Telleria MT, Chaiwan N, Matoéec N, de Silva NI, Pereira OL, Singh PN, Manimohan P, Uniyal P, Shang QJ, Bhatt RP, Perera RH, Alvarenga RLM, Nogal-Prata S, Singh

Luna-Fontalvo et al. | New records of Agaricaceae from Colombia

SK, Vadthanarat S, Oh SY, Huang SK, Rana S, Konta S, Paloi S, Jayasir1 SC, Jeon SJ, Mehmood T, Gibertoni TB, Nguyen TTT, Singh U, Thiyagaraja V, Sarma VV, Dong W, Yu XD, Lu YZ, Lim YW, Chen Y, Tkaléec Z, Zhang ZF, Luo ZL, Daranagama DA, Thambugala KM, Tibpromma S, Camporesi E, Bulgakov TS, Dissanayake AJ, Senanayake IC, Dai DQ, Tang LZ, Khan S, Zhang H, Promputtha I, Cai L, Chomnunti P, Zhao RL, Lumyong S, Boonmee S, Wen TC, Mortimer PE, Xu J (2019) Fungal diver- sity notes 929-1035: taxonomic and phylogenetic contributions on genera and species of fungi. Fungal Diversity 95:1—-273. https:// doi.org/10.1007/s13225-019-00421-w

Pineda F, Garcia G, Velasquez LF, Saldarriaga Y (1988) Descripcion y nuevos registros de Marasmius (Thicholamataceae) en Colombia. Actualidades Bioldgicas 17 (64): 99-106.

Pulido MM (1983) Estudios en Agaricales Colombianos — Los hon- gos de Colombia IX. Instituto de Ciencias Naturales, Museo de Historia Natural, Biblioteca José Jeronimo Triana. Bogota, Co- lombia, 143 pp.

Putzke J, Mejia LGH, Cafion E, Fernandez, Yeina M, Bedoya T (2020) New citations to the Agaricobiota (Fungi - Basidiomycota) in oak forests of the Northeastern Andes of Colombia. Hoehnea 47: 1-9. https://doi.org/10.1590/2236-8906-42/2019

Rangel JO (2015) La biodiversidad de Colombia: significado y dis- tribucion regional. Revista de la Academia Colombiana de Cien- cias Exactas, Fisicas y Naturales 39 (151): 176-200. https://doi. org/10.18257/issn.0370-3908

Rangel-Ch JO, Carvajal-Cogollo JE (2012) Clima de la region Caribe colombiana. In: Rangel-Ch JO (Eds.) Colombia diversidad bidtica XII: la region Caribe de Colombia. Instituto de Ciencias Natura- les, Universidad Nacional de Colombia, Bogota, Colombia, 67— 129,

Rother MS, Da Silveira RMB (2009a) Leucoagaricus lilaceus (Agari- caceae), a poorly known Neotropical agaric. Mycotaxon 107: 473— 487. https://doi.org/10.5248/107.473

Rother MS, Da Silveira RMB (2009b) Leucocoprinus Pat. (Agari- caceae, Basidiomycota) no Parque Estadual de Itapua, Viamao, RS, Brasil. Acta Botanica Brasilica 23 (3): 720— 728. https://doi. org/10.1590/S0102-33062009000300011

Sanchez IM (2003) Composicion de hongos Agaricales en dos bosques de la cuenca de Puerto Abeja, Parque Nacional Natural Chiribi- quete, Caqueta. Bachelor’s degree dissertation, Universidad de los Andes, Bogota, Colombia, 79 pp.

Singer R (1986) The Agaricales in modern taxonomy. Koeltz Scientific Books, Koenigstein, Germany, 981 pp.

Singer R, Digilio APL (1951) Prédromo de la Flora Agaricina Argen- tina. Lilloa 25: 5—461.

Terriquez-Villanueva AK, Herrera-Fonseca MJ, Rodriguez-Alcantar O (2017) Contribucion al conocimiento de la micobiota del cerro Punta Grande, Mezcala, municipio de Poncitlan, Jalisco, México. Revista Mexicana de Micologia 45: 53-66.

Vasco-Palacios AM, Franco Molano AE (2021) Diversity of Co- lombian macrofungi (Ascomycota - Basidiomycota). Version 1.2. Universidad de Antioquia. Dataset/Checklist. https://doi.

1393

org/10.15472/08vo029

Vasco-Palacios, A, Franco-Molano AE (2013) Diversity of Colom- bian macrofungi (Ascomycota-Basidiomycota). Mycotaxon 121 (1): 100-158.

Vellinga EC (2001) Leucocoprinus Pat. In: Noordeloos ME, Kuyper THW, Vellinga EC (Eds.) Flora Agaricina Neerlandica: critical monographs on families of agarics and boleti occurring in the Neth- erlands. A.A. Balkema Publishers, Lisse, Netherlands, 5: 76-84.

Vellinga EC (2003) Chlorophyllum and Macrolepiota (A garicaceae) in Australia. Australian Systematic Botany 16: 361-370. https://do1. org/10.1071/SB02013

Vellinga EC (2006) Lepiotaceous fungi from California, USA — 3. Pink and lilac species in Leucoagaricus sect. Piloselli. Myco- taxon 98: 213-224.

Vellinga EC, Contu E, Vizzini A (2010) Leucoagaricus decipiens and L. erythrophaeus, a new species pair in sect. Piloselli. Mycologia 102: 447-454.

Verma RK, Vimal P, Asati HL (2018) Diversity of macro-fungi in Central India — XI: Leucoagaricus rubrotinctus. Van Sangyan 5 (4): 1-11.

Wijayawardene NN, Hyde KD, Al-Ani LKT, Tedersoo L, Haelewa- ters D, Rajeshkumar KC, Zhao RL, Aptroot A, Leontyev DV, Saxena RK, Tokarev YS, Dai DQ, Letcher PM, Stephenson SL, Ertz D, Lumbsch HT, Kukwa M, Issi IV, Madrid H, Phil- lips AJL, Selbmann L, Pfliegler WP, Horvath E, Bensch K, Kirk PM, Kolafrikova K, Raja HA, Radek R, Papp V, Dima V, Ma J, Malosso E, Takamatsu S, Rambold G, Gannibal PB, Triebel D, Gautam AK, Avasthi S, Suetrong S, Timdal E, Fryar SC, Del- gado G, Réblova M, Doilom M, Dolatabadi S, Pawtowska J, Hum- ber RA, Kodsueb R, Sanchez-Castro I, Goto BT, Silva DKA, de Souza FA, Oehl F, da Silva GA, Silva IR, Blaszkowski J, Jobim K, Maia LC, Barbosa FR, Fiuza PO, Divakar PK, Shenoy BD, Casta- fieda-Ruiz RF, Somrithipol S, Lateef AA, Karunarathna SC, Tib- promma S, Mortimer PE, Wanasinghe DN, Phookamsak R, Xu J, Wang Y, Tian F, Alvarado P, Li DW, Ku8an I, Matoéec N, Ma- harachchikumbura SSN, Papizadeh M, Heredia G, Wartchow F, Bakhshi M, Boehm E, Youssef N, Hustad VP, Lawrey JD, Santi- ago ALCMA, Bezerra JDP, Souza-Motta CM, Firmino AL, Tian Q, Houbraken J, Hongsanan S, Tanaka K, Dissanayake AJ, Mon- teiro JS, Grossart HP, Suija A, Weerakoon G, Etayo J, Tsurykau A, Vazquez V, Mungai P, Damm U, Li QR, Zhang H, Boonmee S, Lu YZ, Becerra AG, Kendrick B, Brearley FQ, Motiejinaite J, Sharma B, Khare R, Gaikwad S, Wijesundara DSA, Tang LZ, He MQ, Flakus A, Rodriguez-Flakus P, Zhurbenko MP, McKenzie EHC, Stadler M, Bhat DJ, Liu Jk, Raza M, Jeewon R, Nasson- ova ES, Prieto M, Jayalal RGU, Erdogdu M, Yurkov A, Schnit- tler M, Shchepin ON, Novozhilov YK, Silva-Filho AGS, Liu P, Cavender JC, Kang Y, Mohammad S, Zhang LF, Xu RF, Li YM, Dayarathne MC, Ekanayaka AH, Wen TC, Deng CY, Pereira OL, Navathe S, Hawksworth DL, Fan XL, Dissanayake LS, Kuhnert E, Grossart HP, Thines M (2020) Outline of Fungi and fungus- like taxa. Mycosphere 11 (1): 1060-1456. https://do1.org/10.5943/ mycosphere/11/1/8