taxonID	type	description	language	source
9A684B45300A7133FBEB7A9AE3A2E61C.taxon	description	Singer (1976: 217) grouped M. ferrugineus and its variety (elevated herein to species), and M. anomalus, M. bambusinus Fr., M. guzmanianus Singer, M. hypophaeus, M. montagneanus Singer, M. nogalesii Singer, M. phaeocystis Singer, and M. tenuisetulosus (Singer) Singer in this stirps. The clade Ferrugineus (Fig. 1) seems consistent with stirps Ferrugineus as it includes M. ferrugineus sensu Antonín et al. (2012) from South Korea with additional strains from China and M. gardneri from Brazil. Data including M. ferrugineus topotypic collections along with other species are needed to further elucidate this group. Based on morphology, M. bambusinus described below seems to belong to this group but sequencing failed in this study. On the other hand, M. anomalus grouped with M. hinnuleus sensu Grace et al. (2019), M. grandisetulosus (very similar to M. tenuisetulosus) and M. hypophaeus sensu Wannathes et al. (2009) forming a monophyletic group distant from clade Ferrugineus. The authentic M. hypophaeus should branch in clade Ferrigineus though.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300A7131FF1A7A3AE692E185.taxon	description	Basidiomata marasmioid (umbrella-like), thin, small- to medium sized. Pileus membranous, sulcate. Lamellae distant to subdistant, free to adnate. Stipe mostly filiform with a scanty, tomentose basal mycelium. Basidiospores elongate, clavate to subfusoid (11 - 25 µm long, xm = 17.5 - 22 µm, Qm = 4.3 - 5.9). Pleurocystidia present, well-developed, elongate, refractive. Pileipellis composed of Siccus-type broom cells only. Habit preference for leafy and / or woody substrate.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300A7131FF1A7A3AE692E185.taxon	materials_examined	TYPE SPECIES. — Marasmius haematocephalus (Mont.) Fr. NOTES In Oliveira et al. (2020), this series included stirpes Ferrugineus, Haematocephalus and Siccus of Singer (1976). These informal supraspecific groups are presented below with morphologically described species (except stirps Siccus) included in the phylogenetic analyses in this study.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530087135FEF17DF4E055E05D.taxon	description	(Figs 6 A; 7)	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530087135FEF17DF4E055E05D.taxon	materials_examined	EXAMINED MATERIAL. — Brazil. São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, 3. III. 2011, J. J. S. Oliveira & F. Karstedt JO 343 (SP [SP 445503]!).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530087135FEF17DF4E055E05D.taxon	biology_ecology	HABIT AND SUBSTRATE. — Marasmioid (Fig. 6 A), gregarious, on a mix of dead sticks, small twigs, tendril and leaves of eudicotyledonous plant or petioles and culm-like sticks in the forest litter.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530087135FEF17DF4E055E05D.taxon	distribution	DISTRIBUTION. — Marasmius bambusinus was originally described from Brazil as Agaricus bambusinus Fr. (Fries 1830), later combined in Marasmius (Fries 1838). It was reported again from Brazil (Pernambuco State) and also from Bolivia, Colombia and Venezuela (Singer 1976). This is the first record from the São Paulo State.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530087135FEF17DF4E055E05D.taxon	description	DESCRIPTION Pileus (Figs 6 A; 7 A) 2 - 13 mm diam., mostly hemispheric to convex, or campanulate, shallowly or deeply sulcate, center flat or slightly depressed, margin decurved, edge entire or slightly crenate, with rare lacerations; brightly orange to fulvous (N 20 Y 80 - 99 M 50 - 60), also pallescent orange (N 00 Y 40 - 60 M 20 - 30), center deep orange to brownish, ferrugineus orange (N 50 Y 99 M 60); membranous, context thin (<1 mm); glabrous, semi humid to dry, dull, subvelutinous, non-hygrophanous. Lamellae (Figs 6 A; 7 A) Free to mainly adnexed, distant, L = 8 - 9, equal, simple, l = 0, opaque, smooth, pale cream (N 00 Y 10 M 10), edge even, nonmarginate, interlamellar hymenium concolorous with the lamellae faces or partly with the pileus. Stipe (Figs 6 A; 7 A) 7 - 22 × 0.2 - 0.4 mm, central, often curved, filiform, thin, equal, with circular caliber, chitinous, flexible, hollow; apex concolorous with the lamellae faces, then becoming pale ochraceous (N 40 Y 99 M 50 to N 50 Y 99 M 50) to dark brown downwards, glabrous, smooth, with a silky bright; with a scanty, off-white, tomentose basal mycelium. Odor Not distinctive. Basidiospores (Fig. 7 B) 16 - 21 × 3.5 - 4 µm (xm = 17.8 [± 1.3] × 3.8 [± 0.1] µm, Qm = 4.7 [± 0.3], n / s = 30 / 1), oblong, clavate to rarely subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia (Fig. 7 C) 23 - 34.4 × 7.3 - 8.4 µm, clavate, smooth, hyaline, 4 - sterigmate, thin-walled, inamyloid. Basidioles (Fig. 7 D) 20 - 25 × 5.3 - 8 µm, similar to the basidia. Pleurocystidia (Fig. 7 E) 43.8 - 56.3 (- 70) × 10 - 16 µm, conspicuous, sometimes with base deepened in the subhymenium, clavate, some slightly capitate or solely with shallow constriction near the apex, smooth, semi translucent or fuscous, refractive, inamyloid, thin-walled at the apex, many times moderately thick-walled elsewhere. Cheilocystidia (Fig. 7 F) Similar to the Siccus-type broom cells of the pileipellis; main body 11.3 - 21.3 × 5 - 8.8 µm, clavate to slightly turbinate, thinwalled, hyaline; setulae or diverticula apical, erect, short to somewhat long, 1.3 - 4.4 × 0.8 - 1 µm, digitiform, cylindrical or verruciform, hyaline, simple, abundant, solid, pale yellow, apex obtuse to slightly acute. Lamellar trama Dextrinoid, irregular, interwoven, hyphae cylindrical, 2 - 8.8 µm diam., regular in outline, branched, hyaline, smooth, thinwalled. Pileus trama Dextrinoid, irregular, very narrow, similar to the lamellar trama, hyphae 1.3 - 5 µm diam. Pileipellis Hymeniform, composed of Siccus-type broom cells (Fig. 7 G), pale yellow when grouped, easily bleaching in KOH solution, becoming hyaline when isolated, abundant; main body 10 - 20 × 6 - 11.3 µm, clavate to slightly turbinate, hyaline, thinwalled; setulae apical, erect, frequently short to moderately long, 2 - 5.6 × 0.5 - 1 µm, digitiform or cylindrical, simple, solid and regular in outline, initially pale yellow, then hyaline, apex acute to almost obtuse. Stipe trama Dextrinoid, cortical hyphae parallel, cylindrical, 3.4 - 7.5 µm diam., regular in outline, smooth, thick-walled, brown to yellowish brown in KOH solution; internal hyphae 1.6 - 8.8 µm diam., thin-walled, some disorganized, other parallel, smooth, branched. Clamp connections Present in almost all tissues, except in the cortical trama of the stipe.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530087135FEF17DF4E055E05D.taxon	discussion	REMARKS Desjardin et al. (2000) considered M. bambusinus a synonym of either M. hypophaeus or M. ferrugineus. These three species have many overlapping morphological characteristics and there is variation of species concepts inDesjardin et al. (2000) and Singer (1976). Basidiospores in these species are quite compatible, but the pigmented lamellar edge (marginate) and the chestnut yellow to fulvous pileus was considered typical and distinctive for M. hypophaeus (Singer 1976). Marasmius bambusinus differs from M. ferrugineus (lignicolous) by a supposed substrate preference on monocotyledonous plant (originally on bamboo leaves), by thin- to thick-walled pleurocystidia, and by a bright orange (- rufescent) pileus (Singer 1976). The holotype of M. bambusinus has been reported as lost (Singer 1976; Desjardin 1989). The examined material brings emphasis for the bright orange (rufescent) pileus (Fig. 6 A) and the more distant, few (paucis) lamellae (8 - 9), both consistent with the protologue of M. bambusinus. Agaricus (Marasmius) ferrugineus Berk. (synonym of Marasmius ferrugineus) was described having a yellow, ferruginous (croceo-ferrugineo) pileus and few lamellae (Berkeley 1843). In Singer (1976), M. ferrugineus has 9 - 13 lamellae. If M. bambusinus (not well-known) is synonym of M. ferrugineus (more well known), then the former has nomenclatural priority. Based onSinger (1976), the examined material can be either M. bambusinus or M. ferrugineus, unless we consider strong enough the fine pileus color deviation and the paucity of the lamellae. Antonín et al. (2012) revised the type of M. ferrugineus and found only slightly larger (overlapping) basidiospores (18 - 22 × 4.5 - 6.0 µm) but compatible pleurocystidia ([28 -] 35 - 50 [- 66] × 11 - 17 µm). Desjardin (1989) studied one of Singer’s M. bambusinus collections (Singer B 6345) from Colombia. It is similar to JO 343 but differs in having reddish lamellar edges (typical for M. hypophaeus), by growing on gramineous leaves, by having smaller basidiospores (15.2 - 18.4 × 3.6 - 4.6 µm), and by having thin-walled, slightly smaller pleurocystidia (30 - 45 × 6.5 - 9 µm). These spores and pleurocystidia are compatible to M. gardneri though (next taxon). Antonín et al. (2012) studied the holotype of M. ferrugineus to support the identification of the foliicolous specimen (BRNM 724480) from South Korea as M. ferrugineus. If so, there is no preference for woody substrate in M. ferrugineus. The lamellae edge concolorous with the brownish orange to reddish brown pileus is rather more consistent with M. hypophaeus (Singer 1976). Without sequences from JO 343, M. bambusinus could not be analyzed along with M. ferrugineus from South Korea (Antonín et al. 2012). Pegler (1988) mentioned M. ferrugineus as a common Neotropical and pantropical species. More should be done to elucidate between M. bambusinus and M. ferrugineus, especially in sequencing collections from the Neotropics (particularly Brazil) matching the morphology and substrate. The species are then regarded as distinct herein. Marasmius pseudobambusinus Desjardin is similar to M. bambusinus, but differs in having indistinctly sulcate pileus, smaller basidiospores (13.6 - 19.2 × 3.6 - 5.2 µm) and apically constricted pleurocystidia (Desjardin 1991).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300E7129FED57B34E33FE0A5.taxon	description	(Figs 6 B; 8)	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300E7129FED57B34E33FE0A5.taxon	materials_examined	EPITYPE. — Brazil. São Paulo State, São Paulo City, Parque Estadual da Cantareira, Núcleo Engordador, 16. II. 2012, J. J. S. Oliveira & M. Capelari JO 491 (epi-, designated here, SP [SP 445564]!). ADDITIONAL EXAMINED MATERIAL. — Brazil. São Paulo State, São Paulo City, Parque Estadual da Cantareira, Núcleo Engordador, 31. X. 2011, J. J. S. Oliveira & M. Capelari JO 387 (SP [SP 445521]!); 19. XII. 2011, J. J. S. Oliveira & M. Capelari JO 438 (SP [SP 445532]!); J. J. S. Oliveira & M. Capelari JO 454 (SP [SP 445542]!).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300E7129FED57B34E33FE0A5.taxon	biology_ecology	HABIT AND SUBSTRATE. — Marasmioid (Figs 6 B; 8 A 1), gregarious, on eudicotyledonous dried leaves and twigs in the forest litter.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300E7129FED57B34E33FE0A5.taxon	distribution	DISTRIBUTION. — This species was established with a nomen novum in Singer (1958) based on the type material of M. ferrugineus originally from Minas Gerais State (Brazil) collected by Gardner. This type consisted of a mix of two close species, later split into two sets where the “ Hooker Herbarium set ” of basidiomata is the type of this species. Additional collections listed by Singer (1958, 1976) are from the Amazonas and Rio de Janeiro States (Brazil).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300E7129FED57B34E33FE0A5.taxon	description	DESCRIPTION Pileus (Figs 6 B; 8 A 1) Up to 2 mm diam. when immature, then 3 - 18 mm diam., conical to campanulate (because of a broad umbo), or hemispheric, becoming convex, sometimes applanate, smooth when young, often becoming slightly sulcate, or strongly sulcate when fully developed, center umbonate, later tending to flat and sometimes wrinkled, margin decurved to straight, edge mostly entire, or slightly crenate; center dark reddish brown (N 70 - 90 Y 90 - 99 M 80 - 90) to dark orangish brown (N 60 Y 99 M 70), then becoming more pale chestnut red or fulvous “ tawny ” (N 60 Y 60 M 60) or dark brown (N 60 Y 70 - 80 M 90 - 99) especially when young, or rarely pale orange (N 10 Y 99 M 30) or pale pinkish brown (N 60 Y 80 M 70), many times with darker sulci, some strongly ferruginous (N 80 Y 99 M 70), orangish brown to fulvous-ferruginous (N 40 - 50 Y 99 M 60 to N 40 - 50 Y 99 M 70) toward the margin, and orange to ferruginous brown when dried (N 80 Y 99 M 50 - 70); membranous, context white, thin (<1 mm); glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae (Figs 6 B; 8 A 2) Free to subfree, subdistant, L = 12 - 16, equal, mostly simple, or very slightly intervenose in very mature basidiomata, l = 0 (- 1), straight to slightly ventricose, opaque, smooth, white, or pale cream to whitish pink (N 00 Y 10 M 00 - 10), edges even, non-marginate, concolorous with the lamellae faces as well as the interlamellar hymenium. Stipe (Figs 6 B; 8 A 1) 15 - 44 × 0.3 - 1 mm, central, filiform, or thicker, almost cylindrical thin, equal, with circular to slightly compressed caliber, simple, chitinous, hollow, apex whitish (N 20 Y 10 M 30) or pale brown (N 30 Y 40 M 30), becoming brown (N 90 Y 90 M 50 - 60), reaching dark brown (N 99 Y 99 M 80) at the base, glabrous, smooth, with a silky bright; with a more abundant, whitish, tomentose or cotton-like basal mycelium, also developing a mat on the substrate. Odor Not distinctive. Basidiospores (Fig. 8 B) (14.6 -) 15 - 19 × 3 - 5 µm (xrm = 16.6 - 17.6 × 3.9 - 4.1 µm; xmm = 16.7 [± 0.6] × 3.9 [± 0.1] µm; Qmr = 4.1 - 4.6, Qmm = 4.3 [± 0.3]; n / s = 30 / 3), oblong, clavate to fusoid, smooth, hyaline, thin-walled, inamyloid. Basidia Not observed. Basidioles (Fig. 8 C) 21.3 - 33 × 5 - 6.3 µm, clavate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia (Fig. 8 D) 32.5 - 62.5 × 4 - 7.5 µm, conspicuous, well-projecting above the basidioles, clavate to cylindrical, apex obtuse or sometimes fusoid to acuminate, or capitate, with a small apical vesicle, or mucronate, smooth, refractive, inamyloid, few fuscous, thin-walled. Cheilocystidia (Fig. 8 E) Similar to the Siccus-type broom cells of the pileipellis, but non-pigmented; main body 12.5 - 23.8 × 5.6 - 8.8 (- 10) µm, clavate to somewhat turbinate, or ventricose, regular in outline, thin-walled; setulae apical, erect, generally elongate, 2.5 - 10.4 × 0.6 - 1.3 µm, cylindrical, filiform, rarely digitiform, regular in outline, simple, pale yellow, solid, apex obtuse to acute. Lamellar trama Dextrinoid, irregular, interwoven, hyphae cylindrical, 2 - 8.8 µm diam., regular in outline, branched, smooth, hyaline, thinwalled. Pileus trama Similar to the lamellar trama, hyphae 1.5 - 8.8 µm diam. Pileipellis Hymeniform, dextrinoid, composed of Siccus-type broom cells (Fig. 8 F), abundant, pale orangish brown when grouped, hyaline when isolated; main body (8.8 -) 12.5 - 18.8 × 5 - 9.4 µm, clavate to turbinate, sometimes cylindrical, or branched, or somewhat irregular in outline, thin- to thick-walled, weakly dextrinoid; setulae apical, erect, 3 - 8.8 × 0.5 - 1.3 µm, cylindrical, filiform, thin, needle-like, simple, rarely branched, pale brown, apex acute or slightly obtuse. Stipe trama Dextrinoid, cortical hyphae parallel, cylindrical, 3.8 - 9.4 µm diam., regular in outline, pale brown, hyaline at the stipe apex, dark brown near the base, smooth, thick-walled; internal hyphae more hyaline, thin-walled, 2 - 16.3 µm diam. Clamp connections Present in almost all tissues, except in the cortical hyphae of the stipe.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45300E7129FED57B34E33FE0A5.taxon	discussion	REMARKS The historical account for this species is not straightforward and it begins with Agaricus (Marasmius) ferrugineus (Berkeley 1843). Due to competing prior homonym, it was later combined by Berkeley & Curtis (1869) in Marasmius ferrugineus. Singer (1958), by arguing that this name was illegitimate in the basionym, proposed the nomen novum Marasmius gardneri in replacement, based on the type material of M. ferrugineus along with two additional collections: 1) ‘ part’ of Spruce 139 (from Amazonas State, Brazil) segregated (mixed collection) from the type of M. poecilus Berk.; and 2) Singer B 437 (from Angra dos Reis, Rio de Janeiro State). However, M. ferrugineus is deemed legitimate in Berkeley & Curtis (1869) and, therefore, M. gardneri became a nom. illegit. by the art. 52.1 of the International Code of Nomenclature for Algae, Fungi and Plants. According to Singer (1976), the type collection of M. ferrugineus was potentially a mix of two different species as the Gardner collection existed split into two sets (Singer 1958), both collected at Minas Gerais and kept at K. The first set, kept in the Berkeley Herbarium, was indicated as the holotype of M. ferrugineus, and has relatively larger and narrower spores (15 - 21 × 2.8 - 4.3 m). The second set, kept in the Hooker Herbarium, has shorter and broader spores (13.5 - 18.3 × 3.2 - 5 m), and Singer (1976) named this collection M. ferrugineus var. gardneri Singer. This variety also differs from the type variety in the pileus often becoming somewhat darker (“ cocoa ”) on drying and in growing on small woody sticks, leaf petioles and veins (Singer 1976). The examined specimens fit best as M. ferrugineus var. gardneri. The dimensions of the pileus (3 - 18 mm diam.) and of the stipe (15 - 44 × 0.3 - 1 mm) are larger than those typically found in M. ferrugineus var. ferrugineus (3 - 11 mm diam. of the pileus and 11 - 30 × 0.3 mm of the stipe). Singer (1976) did not mention the dimensions of the pileus and stipe as distinctive for the specimens of var. gardneri, which implies that both varieties match relatively the macroscopic proportions. The protologue of Agaricus (Marasmius) ferrugineus (synonym of M. ferrugineus) emphasizes the tiny proportion of the basidiomata with “ pileus 1 ½ - 3 lines broad ” (3.18 - 6.35 mm diam.) and stipe “ ½ - ¾ of an inch high, ⅛ of a line thick ” (12.7 - 19.05 mm long and 0.265 mm thick). The basidiomata of the examined material are evidently larger and more robust (as in M. hypophaeus) than a typical M. ferrugineus and, in combination with the shorter and broader basidiospores and by growing more on dried leaves or small twigs, should represent a different species. According to Singer (1958, 1976), Dennis also knew very well M. ferrugineus, especially the Berkeley Herbarium set which, agreeing with Singer, defended it should be the type of M. ferrugineus. With collections from Venezuala, Dennis (1961) found two spore’s dimension for “ M. ferrugineus ”: Dennis 1021 (19 - 21 × 3 - 4 µm) and Dennis 1021 A (14 - 18 × 3 - 3.5 µm, this should be M. gardneri). Revising the type specimen of M. ferrugineus (Brazil, Minas Gerais, K [M] 92652), Antonín et al. (2012) found even longer and broader basidiospores 18 - 22 × 4.5 - 6.0 µm and broader pleurocystidia (11 - 17 µm) which are not consistent with the specimens herein determined as M. gardneri. This is sister to M. ferrugineus from South Korea and China (Fig. 1). Based on the combined divergences found between the two sets of Gardner collection together with the examined collections agreeing with the Hooker Herbarium set, we conclude that M. ferrugineus var. gardneri should be elevated to species level, legitimating the name M. gardneri typified on the ‘ Hooker Herbarium set’. Moreover, if M. ferrugineus becomes a synonym of M. bambusinus (previous taxon), then we are quite sure we have two different species. Both Singer (1958) and Dennis (1951) considered M. paucifolius Murrill, a synonym of M. ferrugineus. Singer (1976) rather placed it in M. ferrugineus var. gardneri and, therefore, should claim its legitimacy for being prior than M. gardneri. However, we have scarce evidence to support M. paucifolius as synonym of M. ferrugineus var. gardneri and should be considered an independent species until more collections and analyses become available. If conspecific, the species would also occur in Puerto Rico. Marasmius hypophaeus is very similar to M. gardneri but differs mainly in having distinctly marginate lamellae (rust brown) and larger basidiospores ([12 -] 14.5 - 21.5 × 3 - 5.5 µm), broader pleurocystidia (6 - 13 µm) and larger setulae (1 - 14 [- 20] × 0.7 - 2.2 µm) on the broom cells of the pileipellis (Singer 1976). Singer (1976) argued that Dennis (1951) described and illustrated a specimen named after M. ferrugineus orangish brown in both pileus and lamellar edge, then he suspected that M. ferrugineus sensu Dennis (1951) would be rather M. hypophaeus. Singer concluded that the pileus pigmentation of M. hypophaeus when fresh would be orangish brown (old bronze) rather than red (rufous blood red) as described in the protologue (Berkeley & Curtis 1869) that possibly led Murrill (1915) to suppose it would be a synonym of M. haematocephalus. Next, Singer defended that M. hypophaeus is closer to M. ferrugineus than M. haematocephalus but differing from the former by the distinctly ferruginous brown lamellae edge instead of concolorous with the lamellae face. Marasmius tenuisetulosus (Singer) Singer is similar to M. gardneri in many features, especially the basidiospores size (14.5 - 19 × 3 - 4 µm). However, the former differs by having larger (up to 28 mm diam.), orangish brown pileus which is radially striped, by having series of lamellulae, and by having thick-walled pleurocystidia (Singer 1964, 1976). These pleurocystidia are long-acuminate and ventricose, tapered from the middle to the apex (Singer 1964). Marasmius radiatus Desjardin is similar to M. gardneri especially in the orange or brownish orange pileus, in the basidiospores size (15.7 - 19.2 × 3.8 - 5.1 µm) and in the general aspects of the pleurocystidia. However, M. radiatus differs only in having a ferruginous pileus with pale radial stripes over the lamellae line when dried, and in having dimorphic, hyaline cheilocystidia: 1) Siccus-type broom cells; and 2) nonsetulose cystidia similar to the pleurocystidia (Desjardin et al. 1992).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530127129FC157CBFE634E6A8.taxon	description	Singer (1976: 208) included M. haematocephalus and all its varieties, plus M. pallescens and M. panerythrus Singer in this group. In this paper, stirps Haematocephalus is monophylectic in haemat _ cp 1, now represented by the M. haematocephalus complex with pantropical distribution (Fig. 1). Even though similar to M. haematocephalus in the pileus color, both M. pallescens and M. panerythrus seem rather morphologically closer to M. pulcherripes (ser. Pulcherripes) by the shorter basidiospores.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453012712BFCB97E7EE77DE0A5.taxon	description	(Figs 9 A; 10) Differs from M. haematocephalus by having pale orange pileus and slightly larger basidiospores (xmm = 21.1 [± 0.7] × 3.8 [± 0.3] µm, Qmm = 5.7 [± 0.5] vs xmm = 20.5 [± 0.3] × 3.9 [± 0.2] µm, Qmm = 5.2 [± 0.1]); and more distinctly larger than those of the rose or pale red piliated M. rubicundus (xmm = 21.1 [± 0.7] × 3.8 [± 0.3] µm, Qmm = 5.7 [± 0.5] vs xmm = 18.9 [± 0.5] × 3.8 [± 0.1] µm, Qmm = 5 [± 0.2]).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453012712BFCB97E7EE77DE0A5.taxon	materials_examined	HOLOTYPE. — Brazil. São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 2. XII. 2009, M. Capelari & L. A. S. Ramos 4554 (holo-, SP [SP 445584]!). ADDITIONAL EXAMINED MATERIAL. — Brazil. São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 16. III. 2010, J. J. S. Oliveira JO 42 (SP [SP 445408]!); 15. X. 2010, J. J. S. Oliveira & C. L. A. Pires JO 224 (SP [SP 445445]!); 16. X. 2010, J. J. S. Oliveira & C. L. A. Pires JO 226 (SP [SP 445446]!); 7. XI. 2010, J. J. S. Oliveira & A. V. Costa JO 276 (SP [SP 445460]!): J. J. S. Oliveira & A. V. Costa JO 277 (SP [SP 445461]!); J. J. S. Oliveira & A. V. Costa JO 279 (SP [SP 445463]!); 7. XII. 2010, J. J. S. Oliveira, A. V. Costa & P. O. Ventura JO 282 (SP [SP 445465]!); J. J. S. Oliveira & P. O. Ventura JO 300 (SP [SP 445479]!).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453012712BFCB97E7EE77DE0A5.taxon	etymology	ETYMOLOGY. — Based on the orange pileus. MYCOBANK. — MB 842535.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453012712BFCB97E7EE77DE0A5.taxon	description	HABIT AND SUBSTRATE. — Marasmioid (Figs 9 A; 10 A), dispersed to gregarious on dead eudicotyledonous leaves in the forest litter. DESCRIPTION Pileus 2.4 - 19 mm diam. (Figs 9 A; 10 A), initially conical, then campanulate or hemispherical, sulcate, sometimes deeply sulcate, center flat or slightly umbonate, wrinkled when dried, margin decurved, edge entire; when young deep reddish orange (N 10 Y 80 M 30 to N 10 Y 99 M 60, N 40 Y 70 M 70), with center dark orange (N 10 Y 99 M 70), becoming yellowish orange or pale orange (N 00 Y 80 M 30, N 10 Y 99 M 50 to N 20 Y 60 M 50) when mature, sometimes with beige hue (N 00 Y 40 - 60 M 20 - 40 to N 10 Y 50 M 50 - 60) or ferruginous orange (N 20 Y 99 M 60) to reddish brown (N 20 Y 50 M 50) when dried, keeping the same pigmentation at the center or becoming chestnut orange (N 30 Y 99 M 50 - 70, N 40 Y 99 M 80 or N 50 Y 99 M 70); membranous, context thin (<1 mm); glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae (Fig. 9 A) Free, subfree to narrowly adnate, subdistant to distant, L = 9 - 14, equal, simple, rarely biforked or intervenose, l = 0, smooth, pale cream (N 00 Y 10 M 00), edge even, non-marginate, interlamellar hymenium concolorous with the lamellae faces. Stipe (Figs 9 A; 10 A) 9 - 56 × 0.3 - 0.8 mm, central, filiform, thin, equal, regular, with circular caliber, chitinous, flexible, hollow, apex whitish pink (N 00 Y M 20 - 60) to pale orange, becoming orange or amber brown (N 40 Y 60 M 50) to brown or dark brown (N 70 Y 99 M 60) toward the base, glabrous, smooth, with a silky bright; with a scarce, cream, tomentose basal mycelium. 10 - 40 Odor Not distinctive. Basidiospores (Fig. 10 B) 17.1 - 24.8 × 2.8 - 5 µm (xrm = 19.4 - 22.3 × 3.5 - 4.2 µm; xmm = 21.1 [± 0.7] × 3.8 [± 0.3] µm; Qrm = 5.1 - 6.3; Qmm = 5.7 [± 0.5], n / s = 30 / 8), exceptional spore size of JO 226 with 17 - 21.9 × 3 - 4.2 (- 5) µm (xm = 19.4 [± 1.0] × 3.6 [± 0.4] µm; Qm = 5.4 [± 0.7]; n / s = 30, s = 1), oblong, subclavate to fusoid, smooth, hyaline, thin-walled, inamyloid. Basidia Observed in JO 279, 20.1 - 30.4 × 6 - 7.6 µm, clavate, hyaline, thin-walled, inamyloid, 4 - sterigmate, sterigmata coarse or broad, with obtuse apex. Basidioles (Fig. 10 C) 21.3 - 28.8 × 6 - 8.8 µm, clavate, hyaline, thin-walled, inamyloid. Pleurocystidia (Fig. 10 D) (15.1 -) 22.3 - 74.3 × 5.2 - 12.1 µm, clavate, with tapered apex, subacuminate, vesiculose, capitate or papillate, sublageniform, hyaline, thin-walled, refractive. Cheilocystidia (Fig. 10 E) Similar to the Siccus-type broom cells of the pileipellis; main body (10 -) 15 - 18.8 × 5.6 - 11.3 µm, clavate to turbinate, hyaline, thin-walled; setulae apical, erect, 2.5 - 7.5 × 0.8 - 1.8 µm, cylindrical, regular in outline, solid, apex acute. Lamellar trama Dextrinoid, irregular, interwoven, hyphae cylindrical, 1.6 - 10 µm diam., regular in outline, branched, smooth, hyaline, thin-walled. Pileus trama Similar to the lamellar trama, hyphae 2 - 6.3 µm diam. Pileipellis Hymeniform, composed of Siccus-type broom cells (Fig. 10 F), abundant, pale yellow when grouped; main body 9.4 - 18.8 × 7 - 10.6 µm, clavate to frequently turbinate, sometimes inflated, or irregular in outline, hyaline, thin-walled, weakly dextrinoid; setulae apical, erect, 3.8 - 7.5 × 0.6 - 1.3 µm, filiform, thin, regular in outline, sometimes branched, solid, hyaline, apex acute. Stipe trama Dextrinoid, cortical hyphae parallel, 3 - 13.8 µm diam., cylindrical, regular in outline, rarely branched, smooth, pale brown, thick-walled; internal hyphae 2.5 - 7.5 µm diam., hyaline, thin-walled. Clamp connections Present in all tissues. REMARKS Marasmius auranticapitatus J. S. Oliveira, sp. nov. is very similar to M. haematocephalus (blood red pileus) but has more pale yellowish orange pileus (reddish when young) and slightly longer basidiospores (xmm = 21.1 [± 0.7] × 3.8 [± 0.3] µm, Qmm = 5.7 [± 0.5] vs xmm = 20.4 [± 0.3] × 3.9 [± 0.1] µm, Qmm = 5.3 [± 0.1]). Yet, the ranges of the basidiospores sizes are largely overlapping. Marasmius auranticapitatus J. S. Oliveira, sp. nov. is 0.2 - 1 % dissimilar to M. haematocephalus based on the nrITS and 0.6 - 1 % dissimilar on the nrLSU. However, M. auranticapitatus J. S. Oliveira, sp. nov. is closer and sister to M. rubicundus (Singer) J. S. Oliveira, stat. nov. (Figs 4; 5). Based on the nrITS data, M. auranticapitatus J. S. Oliveira, sp. nov. is 0.7 - 1.7 % dissimilar to M. rubicundus (Singer) J. S. Oliveira, stat. nov., 0.2 - 0.6 % dissimilar in the nrLSU, 4.7 - 5.2 % dissimilar in the rpb 2 and 1.4 - 1.9 dissimilar in the ef 1 - α. Marasmius auranticapitatus J. S. Oliveira, sp. nov. differs from M. rubicundus (Singer) J. S. Oliveira, stat. nov. in the pale yellowish orange pileus (vs rose, pink or pale red) and longer basidiospores (xmm = 21.1 [± 0.7] × 3.8 [± 0.3] µm, Qmm = 5.7 [± 0.5] vs xmm = 18.9 [± 0.5] × 3.8 [± 0.1] µm, Qmm = 5 [± 0.2]). With a transitional / intermediate pattern and possibly a hydrid, the exceptional spore sizes of JO 226 (more compatible with M. rubicundus (Singer) J. S. Oliveira, stat. nov.) strengthens the plausibility of the most recent common ancestry between M. auranticapitatus J. S. Oliveira, sp. nov. and M. rubicundus (Singer) J. S. Oliveira, stat. nov., a bridge of intercompatibility and gene flow or an ancestry trace. QuotingTaylor et al. (2000): “ Even when BSR (Biological Species Recognition) can be applied to fungi, there is evidence that the criterion of reproduction lumps together groups of fungi that are genetically isolated in nature, but that retain the ancestral character of interbreeding ”. Marasmius auranticapitatus J. S. Oliveira, sp. nov. is similar to M. siccus in the pileus pigmentation. However, based on the holotype, M. siccus has smaller basidiospores 15.2 - 20 × 3.2 - 4.6 µm (Desjardin 1989). Also, M. siccus s. l. has more robust basidiomata (Gilliam 1976; Noordeloos 1987; Antonín & Noordeloos 2010; Antonín et al. 2012;) and temperate distribution. In the phylogenetic trees (Fig. 1), M. auranticapitatus J. S. Oliveira, sp. nov. is close to M. haematocephalus (embedded in haemat _ cp 1) and distant from M. siccus. Marasmius auranticapitatus J. S. Oliveira, sp. nov. is part of the haemat _ cp 2 a (Fig. 3), only fully resolved in Figure 5. Marasmius haematocephalus var. anomaloides Desjardin originally described from the Tennessee, United States, also has orange pileus but differs from M. auranticapitatus J. S. Oliveira, sp. nov. in having much shorter basidiospores (14.4 - 19.2 × 3.4 - 4.6 µm) (Desjardin 1991). The new species is neighter similar to any of the varieties of M. haematocephalus in Singer (1976) nor to any of the heterotypic sysnonyms once listed under M. haematocephalus (Appendix).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	description	(Figs 9 B; 11)	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	diagnosis	Differs from Marasmius davidii Antonín in having non-marginate lamellae, slightly smaller basidiospores (20 - 26 × 3 - 4 µm vs 20 - 27 × 4 - 6 µm), in having yellowish pleurocystidia, and in growing on dried leaves.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	materials_examined	HOLOTYPE. — Brazil. São Paulo State, Iporanga City, Parque Estadual Turístico do Alto Ribeira, 29. II. 2012, J. J. S. Oliveira & D. E. Desjardin JO 523 (holo-, SP [SP 445573]!).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	etymology	ETYMOLOGY. — From Greek Καστανός (kastanos) and ΚΕΦάλι (kephali), based on the brownish pileus.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	description	MYCOBANK. — MB 842536.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	biology_ecology	HABIT AND SUBSTRATE. — Marasmioid (Figs 9 B; 11 A 1), close, on dried eudicotyledonous leaves in the forest litter.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	description	DESCRIPTION Pileus (Figs 9 B; 11 A 1) 7.5 - 15.7 mm diam., hemispheric to conical-campanulate, sulcate, center flat or slightly umbonate, margin decurved, edge entire to crenate; center to mid disc light chestnut to buff brown (N 50 Y 60 - 80 M 30 - 40), margin pale sepia brown (N 40 Y 40 M 30) to pale buff brown or brownish beige (N 20 Y 30 M 20 to N 10 Y 30 - 40 M 10); membranous, context white, thin (<1 mm); glabrous, dry, dull, papyraceous to subvelutinous, non-hygrophanous. Lamellae (Fig. 11 A 2) Free to adnexed, subclose to subdistant, L = 11 - 12, equal, narrow to slighty broad, simple, l = 0, opaque, smooth, pale cream (N 00 Y 10 M 00), edge even, non-marginate, interlamellar hymenium concolorous with the lamellae faces. Stipe (Figs 9 B; 11 A 1) 61 - 80 × 0.5 - 0.8 mm, central, filiform, thin, equal, with circular caliber, chitinous, tough, hollow, apex concolorous with the lamellae, amber (N 60 Y 50 M 30) to dark brown (N 90 Y 70 - 99 M 50), or almost black downwards, glabrous, smooth, with a silky bright; with a scarce, cream, tomentose basal mycelium. Odor Not distinctive. Basidiospores (Fig. 11 B) 20 - 26 × 3 - 4 µm (xm = 22.6 [± 1.2] × 3.7 [± 0.3] µm, Qm = 6.1 [± 0.5], n / s = 32 / 1), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia Not observed. Basidioles (Fig. 11 C) (16.3 -) 20 - 25 × 5 - 8 µm, clavate, rarely tending to acuminate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia (Fig. 11 D) 33 - 50 × 9.8 - 13.8 µm, abundant, broadly clavate, sometimes tapered towards the apex, some capitate, or with a small apical vesicle, or mucronate, smooth, refractive, with faint content, somewhat fuscous, yellowish, thin-walled, inamyloid. Cheilocystidia (Fig. 11 E) Similar to the Siccus-type broom cells in the pileipellis, pale brown when grouped, main body 8 - 22.5 × 5 - 15 µm, clavate to slightly turbinate, or pyriform to almost globose, sometimes ventricose, thin-walled; setulae apical, erect, short to generally elongate, 2.5 - 8.8 × 0.6 - 1 µm, cylindrical, filiform, thin, rarely digitiform, regular in outline, simple, pale yellow to hyaline, solid, apex somewhat obtuse or more frequently acute. Lamellar trama Dextrinoid, irregular, interwoven, hyphae cylindrical, 2.5 - 17.5 µm diam., regular or irregular in outline, branched, hyaline, smooth, thin-walled. Pileus trama Dextrinoid, irregular, hyphae interwoven, cylindrical, 1.3 - 8 µm diam., similar to the lamellar trama, but with some inflated hyphal segments. Pileipellis Hymeniform, composed of Siccus - type broom cells (Fig. 11 F), abundant, yellowish to pale chestnut when grouped; main body 8.8 - 18.8 × 5.6 - 14 µm, clavate to turbinate, rarely globose or branched, hyaline, thin-walled; setulae apical, erect, 2 - 7.5 × 0.5 - 1.3 µm, cylindrical, thin, filiform, simple, regular in outline or contorted, yellowish, solid, apex acute or slightly obtuse. Stipe trama Dextrinoid, cortical hyphae parallel, cylindrical, 1.8 - 8.8 µm diam., regular in outline, branched, brownish, smooth, thick-walled; internal hyphae hyaline, thin-walled, 2.5 - 8 µm diam. Clamp connections Present in all tissues, except in the cortical hyphae of the stipe.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453017712EFF5D7A1AE76FE0A2.taxon	discussion	REMARKS Marasmius castanocephalus J. S. Oliveira, sp. nov. is mainly characterized by the buff brown to brownish beige pileus, the large basidiospores (20 - 26 × 3 - 4 µm) and somewhat fuscous to yellowish pleurocystidia (33 - 50 × 9.8 - 13.8 µm). It seems a member of stirps Helvolus (Singer 1976) in ser. Haematocephali (sensu Singer 1976) because of the brown pileus as in M. helvolus Berk. Marasmius castanocephalus J. S. Oliveira, sp. nov. is closer to M. davidii from Sri Lanka (Indo-Malayan realm), initially described as M. helvolus var. brunneolus Berk. & Broome (Berkeley & Broome 1873). Pegler (1986) elevated this variety to M. brunneolus (Berk. & Broome) Pegler. Marasmius helvolus has much smaller basidiospores (11 - 15 × 2.8 - 4 µm) and not projecting, inconstant, smaller (19 - 25 × 5 - 10.2 µm), opaque and thick-walled pleurocystidia (Singer 1976). Due to the homonym with M. brunneolus (Beeli) Singer, in sect. Globulares (traditional sense) (Singer 1964), Antonín (2003) replaced M. brunneolus (Berk. & Broome) Pegler with M. davidii. This species differs from M. castanocephalus J. S. Oliveira, sp. nov. mainly in having very distant, marginate lamellae (brown edge), slightly larger basidiospores (20 - 27 × 4 - 6 µm), only hyaline pleurocystidia and brownish cheilocystidia (setulae), and basidiomata found on dead sticks (Pegler 1986). Marasmius brunneolus var. fuliginosus Desjardin & E. Horak was described having dark brown to fuliginous pileus (Desjardin & Horak 1997). Marasmius castanocephalus J. S. Oliveira, sp. nov. is close to M. haematocephalus var. leucophyllus Singer, and to M. musicola Murrill, M. allocystis Singer, and M. oleiger Singer (stirps Oleiger) because of the brownish pigmentation of the pileus within ser. Haematocephali sensu Singer (Singer 1976). All these taxa, however, have much shorter basidiospores (up to 20.5 µm) and the last three have distinct pleurocystidia with oil-like content. No other formally published variety under M. haematocephalus seems close. Marasmius semipellucidus Berk. & Broome is discarted as conspecific with M. castanocephalus J. S. Oliveira, sp. nov. (Appendix) by the translucent stipe and geography of the original collection. In the phylogenetic trees (Figs 1; 3), M. castanocephalus J. S. Oliveira, sp. nov. appears close to M. roseus J. S. Oliveira, sp. nov., but with unsupported resolution. Both species branched in haemat _ cp 1. Marasmius castanocephalus J. S. Oliveira, sp. nov. is close to M. haematocephalus but is 4.4 % dissimilar in nrITS and 2.6 % in nrLSU and differs by the pale brown pileus and larger basidiospores.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530147121FF327CBFE374E4B9.taxon	description	(Figs 9 C; 12) Differs from Marasmius asiaticus Mesic & Tkalcec (synonym: M. distantifolius Y. S. Tan & Desjardin) in not having violet or ruby nor bright pink pileus which is rather larger (3.5 - 17 mm diam. vs 2 - 8 mm diam.), in having more numerous (10 - 12) non-marginate lamellae, in having smaller basidiospores (13.8 - 20 × 3.3 - 4.4 µm vs 18.5 - 23 [- 25] × 3.5 - 5 [- 6] µm), in having shorter and especially broader pleurocystidia (35 - 51.3 × 12 - 18.8 µm vs 37 - 42 × 9 - 12 µm), and in having non-mottled pileipellis.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530147121FF327CBFE374E4B9.taxon	materials_examined	HOLOTYPE. — Brazil. São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 9. XII. 2010, J. J. S. Oliveira, P. O. Ventura & A. V. Costa JO 323 (holo-, SP [SP 445491]!). ADDITIONAL MATERIAL EXAMINED. — Brazil. São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, 1. III. 2011, J. J. S. Oliveira & F. Karstedt JO 339 (SP [SP 445502]!).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530147121FF327CBFE374E4B9.taxon	etymology	ETYMOLOGY. — Due to its close relationship as sister to M. asiaticus, in the Tropical Asia. MYCOBANK. — MB 842537.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B4530147121FF327CBFE374E4B9.taxon	description	HABIT AND SUBSTRATE. — Marasmioid (Figs 9 C; 12 A), gregarious, on dried eudicotyledonous leaves and twigs in the forest litter. DESCRIPTION Pileus (Figs 9 C; 12 A) 3.5 - 17 mm diam., hemispheric to convex, becoming plane, slightly sulcate, center flat or with a shallow depression, margin decurved to straight, edge entire to slightly crenate; mostly brownish red (N 60 Y 70 M 90), sometimes brownish pink (N 40 Y 60 M 60 - 70) or orangish red (N 50 Y 80 M 70), with dark reddish brown center (N 80 Y 99 M 80); membranous, context thin (<1 mm); glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae (Figs 9 C; 12 A) Adnate, subdistant to distant, L = 10 - 12, equal, simple or slightly intervenose, l = 0, opaque, cream (N 00 Y 10 M 00), edge even, nonmarginate, interlamellar hymenium concolorous with the pileus. Stipe (Figs 9 C; 12 A) 9 - 20 × 0.3 - 0.5 mm, central, generally short, curved, filiform, thin, equal, with circular caliber, chitinous, hollow, apex concolorous with the lamellae, becoming black elsewhere, glabrous, smooth, glossy; with a scarce, white to cream, tomentose basal mycelium. Odor Not distinctive. Basidiospores (Fig. 12 B) 13.8 - 20 (- 20.6) × 3.3 - 4.4 µm (xmr = 17 - 18.2 × 3.8 - 3.9 µm; xmm = 17.6 [± 0.8] × 3.9 [± 0.1] µm; Qrm = 4.4 - 4.8; Qmm = 4.6 [± 0.3]; n / s = 30 / 2), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia (Fig. 12 C) 19.4 - 25 × 6.6 - 9 µm, broadly clavate, smooth, hyaline, 2 - 4 - sterigmate, inamyloid, thin-walled. Basidioles (Fig. 12 D) 17.3 - 22.5 × 7 - 9.8 µm, broadly clavate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia (Fig. 12 E) (16.3 -) 35 - 51.3 × 12 - 18.8 µm, abundant, broadly clavate to inflated, bacilliform, almost oval, smooth, slightly fuscous, yellowish, sometimes hyaline, thin-walled, refractive, inamyloid. Cheilocystidia (Fig. 12 F) In form of Siccus-type broom cells, hyaline, walls thinner than those of the pileipellis; main body 10 - 16.3 × 5.6 - 11.3 µm, cylindrical to clavate, sometimes pyriform to turbinate, thinwalled, hyaline; setulae apical, sometimes slightly divergent, erect, generally short, 1.3 - 3.8 (- 5) × 0.6 - 1 µm, filiform, cylindrical, digitiform, or conical to verruciform, simple or rarely branched, solid, hyaline, regular to slightly irregular in outline, apex obtuse to somewhat acute. Lamellar trama Dextrinoid, irregular, interwoven, hyphae cylindrical, 2 - 6.3 µm diam., regular or irregular in outline, smooth, hyaline, branched and strongly connected, thin-walled. Pileus trama Similar to the lamellar trama, hyphae 2 - 7.5 µm diam. Pileipellis Hymeniform, composed of Siccus-type broom cells (Fig. 12 G), abundant, brownish when in group; main body 11.3 - 18.8 × 6.3 - 16.3 µm, clavate to turbinate, sometimes flattened, or branched, or irregular in outline, hyaline, thin- to somewhat thick-walled; setulae or diverticula apical, sometimes slightly divergent, erect, 0.6 - 3.8 (- 5) × 0.5 - 1.1 µm, cylindrical, digitiform, or verruciform to vesiculose, simple to branched, regular to irregular in outline, almost monilioid, solid, hyaline, or melleus to pale brown, apex obtuse to slightly acute. Stipe trama Slightly dextrinoid, cortical hyphae parallel, cylindrical, regular in outline, 2.5 - 8.8 µm diam., branched, smooth, dark brown, yellowish brown when separated, thick-walled; internal hyphae hyaline, 1.3 - 11.3 µm diam., thin-walled. Clamp connections Present, except in the cortical hyphae of the stipe. REMARKS Marasmius coasiaticus J. S. Oliveira, sp. nov. is morphologically very close to M. haematocephalus, but produces smaller, thinner and more delicate basidiomata, having often short and curved stipe. The basidiospores are shorter in average (17 - 18.2 µm vs 20.1 - 21 µm) and the pleurocystidia are pale yellow, distinctly broadly clavate or bacilliform inflated (12 - 18.8 µm vs 7 - 9.9 µm in width). Finally, the apical setulae of the Siccus-type broom cells are many times shorter and verruciform. Marasmius coasiaticus J. S. Oliveira, sp. nov. is 1.8 % dissimilar to M. haematocephalus in nrITS. The evident broader pleurocystidia combined with the shorter basidiospores in M. coasiaticus J. S. Oliveira, sp. nov. easily separate it from all varieties under M. haematocephalus (Singer 1976). Marasmius sanguineus Cooke & Massee may be similar, but differs in the adnexed, ventricose, marginate lamellae and longer stipe (up to 40 mm vs up to 20 mm) with lower pileus / stipe length proportion (Cooke 1889; Appendix). Marasmius coasiaticus J. S. Oliveira, sp. nov. from Brazil is rather sister to M. asiaticus from Malaysia and additional collections from India and Thailand (Figs 1; 3) and, therefore, distant from M. auranticapitatus J. S. Oliveira, sp. nov., M. castanocephalus J. S. Oliveira, sp. nov., M. haematocephalus, M. roseus J. S. Oliveira, sp. nov. and M. rubicundus (Singer) J. S. Oliveira, stat. nov. from Brazil. Marasmius asiaticus was originally named M. distantifolius but, due to competing homonymy with Marasmius distantifolius (Murrill) Murrill, was renamed in Mesic & Tkalcec (2010). Marasmius asiaticus (synonym: M. distantifolius) differs from M. coasiaticus J. S. Oliveira, sp. nov. by having ruby to pinkish hues in the brownish red to brownish violet pileus, marginate, 7 - 8 lamellae, larger basidiospores (18.5 - 23 [- 25] × 3.5 - 5 [- 6] µm), smaller pleurocystidia (37 - 42 × 9 - 12 µm), and mottled pileipellis (Tan et al. 2009).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301A7125FF0C787EE377E4C6.taxon	description	(Figs 13; 14)	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301A7125FF0C787EE377E4C6.taxon	materials_examined	EPITYPE. — Brazil. São Paulo State, Iporanga City, Parque Estadual Turístico do Alto Ribeira, Núcleo Santana, 02. III. 2012, J. J. S. Oliveira & D. E. Desjardin JO 533 (epi-, designated here, SP [SP 445580]!), nrITS (ON 502673) and nrLSU (ON 502729). ADDITIONAL EXAMINED MATERIAL. — Brazil. São Paulo State, São Paulo City, Parque Estadual da Cantareira, Núcleo Engordador, 16. II. 2012, J. J. S. Oliveira & M. Capelari JO 507 (SP [SP 446044]!); Iporanga City, Parque Estadual Turístico do Alto Ribeira, Núcleo Santana, 04. XII. 2011, J. J. S. Oliveira JO 435 (SP [SP 446079]!); 29. II. 2012, J. J. S. Oliveira & D. E. Desjardin JO 527 (SP [SP 446068]!); Amazonas State, Novo Airão City, Urubuquara, Rio Unini, 24. X. 2019, J. J. S. Oliveira & Francisco JO 1363 (INPA [INPA 289980]!).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301A7125FF0C787EE377E4C6.taxon	biology_ecology	HABIT AND SUBSTRATE. — Marasmioid (Figs 13; 14 A 1), close, on dried eudicotyledonous petioles in the forest litter.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301A7125FF0C787EE377E4C6.taxon	distribution	DISTRIBUTION. — The species was originally described from Rio de Janeiro, Brazil. Based on the present study, it is only confirmed in the Neotropics. In Brazil, it was reported from Amazonas State (Berkeley 1856; Berkeley & Cooke 1876; Hennings 1904; this present study), Paraná State (de Meijer 2001), Pernambuco State (Kimbrough et al. 1995), Rio de Janeiro State (Singer 1976), Rondônia State (Capelari & Maziero 1988) and São Paulo State (Spegazzini 1889; Grandi et al. 1984; Pegler 1997; Puccinelli & Capelari 2009). However, these collections need to be revised, especially from the other states far from Southeastern Brazil. Based on the traditional view, the species is pantropical, occurring in tropical forests of South America, Africa, Tropical Asia and Oceania.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301A7125FF0C787EE377E4C6.taxon	description	DESCRIPTION Pileus (Figs 13; 14 A 1) 2.5 - 16 mm diam., conical, hemispherical to convex, or campanulate, deeply sulcate, center flat, or wrinkled, margin decurved, edge entire to slightly crenate; dark purplish red (N 90 Y 00 - 40 M 99) when young, then deep to dark blood red (N 70 Y 50 - 90 M 99) or deep purplish red (N 80 Y 10 - 50 M 99), dark pinkish red almost purple (N 60 Y 50 M 80); membranous, context thin (<1 mm); glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae (Figs 13; 14 A 2) Free to narrowly adnate, distant, L = 7 - 10, equal, narrow, subcultriform, simple, l = 0, opaque, smooth, white to cream (N 00 Y 10 M 00), or whitish pink (N 00 Y 10 M 10 or N 00 Y 40 M 10 - 20), edges even, non-marginate, interlamellar hymenium concolorous with the lamellae faces or partly concolorous with the pileus (especially near the pileus edge). Stipe (Figs 13; 14 A 1) 22.4 - 62 × 0.2 - 0.5 mm, central, filiform, thin, equal, sometimes with slightly broader base, with circular caliber, chitinous, hollow; apex concolorous with the lamellae, becoming bronze brown (N 40 Y 60 M 50 to N 80 Y 70 M 40) to dark brown (N 90 Y 99 M 80), or almost black at the base, glabrous, smooth, with a silky bright; with a scarce (subinsititious), white, tomentose basal mycelium. Odor Not distinctive. Basidiospores (Fig. 14 B) (18 -) 18.3 - 22.6 (- 23) × 3 - 4.9 µm (xrm = 20.1 - 21 × 3.8 - 4.2 µm; xmm = 20.5 [± 0.3] × 3.9 [± 0.2] µm; Qrm = 5 - 5.4; Qmm = 5.2 [± 0.1]; n / s = 30 / 5), oblong, clavate, subfusoid to fusoid, smooth, hyaline, thin-walled, inamyloid. Basidia (Fig. 14 C) 19 - 26.8 × (5.6 -) 6.2 - 7.9 µm, clavate, smooth, hyaline to slightly fuscous, thin-walled, with four short and conical obtuse sterigmata, inamyloid. Basidioles (Fig. 14 D) (15 -) 18.5 - 25.9 × (3.3 -) 4.8 - 7.1 µm, cylindrical clavate, clavate, sometimes wavy or with tapered apex, smooth, hyaline to slightly fuscous, inamyloid. Pleurocystidia (Fig. 14 E) 30.1 - 76.2 × 7 - 13.9 (- 15.5) µm, clavate or cylindrical clavate, lageniform, some capitate, papillate or rarely mucronate, occasionally wavy with apical and shallow constrictions, smooth, fuscous, thin-walled, refractive, inamyloid, abundant although sparse. Cheilocystidia (Fig. 14 F) Similar to the Siccus-type broom cells of the pileipellis, but hyaline and with thinner walls; main body 10.3 - 19.3 × 5.5 - 10.6 µm, clavate to slightly turbinate, sometimes flat, seldom branched; setulae apical, erect, 2.1 - 5.2 × 0.5 - 1 µm, cylindrical or filiform, needle-like, rarely digitiform, regular in outline, simple or rarely branched, solid, hyaline to fuscous, apex acute. Lamellar trama Strongly dextrinoid, irregular, interwoven, hyphae cylindrical, 1.8 - 5.4 µm diam., regular in outline, branched, hyaline, thin-walled, smooth to slightly rough. Pileus trama Strongly dextrinoid, similar to the lamellar trama, hyphae 1.8 - 6.2 µm diam. Pileipellis Hymeniform, composed of Siccus-type broom cells (Fig. 14 G), pale brown or chestnut brown, bleaching in KOH solution; main body 7.2 - 20.4 × 5.5 - 12.1 µm, clavate, turbinate, sometimes branched, ventricose, or flat, hyaline, thin-walled to slightly thick-walled (particularly at the apex), inamyloid; setulae apical, erect, 2.8 - 6.2 × 0.6 - 1.3 µm, cylindrical or filiform, needle-like, rarely digitiform, simple, regular in outline, solid, mostly pale brown, apex tapered, obtuse to slightly acute. Stipe trama Dextrinoid, especially the internal hyphae and those of the stipe apex, cortical hyphae parallel, packed, cylindrical, regular in outline, 2.7 - 7.7 µm diam., or slightly inflated, sometimes branched, smooth, dark chestnut brown, yellowish brown when separate, thick-walled; internal hyphae regular in outline, 1.8 - 8.1 µm diam., hyaline, parallel or interwoven, sometimes with short segments. Clamp connections Present in all tissues, except in the cortical hyphae of the stipe.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301A7125FF0C787EE377E4C6.taxon	discussion	REMARKS Marasmius haematocephalus is the type species of Marasmius sect. Sicci subsect. Siccini ser. Haematocephali sensu Singer (1976) which is characterized by the presence of pleurocystidia, otherwise compatible with ser. Leonini (Singer 1976). The species epithet means blood red head, (from Greek αίματΟς [haimatos] and ΚΕΦάλι [kephali]), “ rubro - sanguineo ” pileus in the protologue. According to a broad concept, this species seems to produce basidiomata with a high variability of characteristics, especially in the pileus pigmentation, length and shape of both spores and pleurocystidia, and substrate preference. Based on Singer (1958), M. haematocephalus is typically, as the description in the protologue indicates, uniformly blood red, purple, deep purple (not violet) pileated. In Singer (1965, 1976), the sulcate pilei have a large color range of variation from pink, red to purple, even brownish, and with shapes from campanulate, hemispherical, convex to plane. Even obeying a reasonable morphological species spectrum, the criptic nature of taxa in species complex and / or a non-fine species characterization has made M. haematocephalus a receptacle of many misdetermined specimens with an allegedly pantropical distribution. After matching Montagne (1837) and Fries (1838) concepts, the topotypical examined material also agrees with Singer (1958, 1965, 1976) in nearly all aspects. We intended to examine “ R. Singer C 3172 ” and confirm compatible spores’ dimension. We contacted BAFC herbarium to search for Singer’s topotype collection, but without reply during this pandemic time. Antonín (2007) and Shay et al. (2017) mentioned the collection as neotype but did not examine it. Vladimír Antonín (pers. comm.) informed he has a specimen on loan from PC herbarium named as M. haematocephalus collected by Auguste de Saint Hilaire from Brazil that is possibly the authentic type. He shared pictures of the dried specimen that seems macromorphologically quite simitar to our specimens. The examined material herin also agrees with “ French Guyana, Ann. 1850, M. Leprieur nº 990 (PC) ”, determined by Montagne as M. haematocephalus, and fully revised by Desjardin (1989), but the basidiospores in the Leprieur’s collection seems smaller (16 - 21.6 × 4.4 - 5.6 µm, xm = 19.1 [± 1.3] × 4.7 [± 0.4] µm, Q = 3.5 - 4.6, Qm = 4.1 [± 0.3], n = 18). There was a probable confusion in Dennis (1951); where M. tageticolor Berk., should be M. haematocephalus from Trinidad (fresh collection), Dominique (type of M. sanguineus Cooke & Massee) and Bahamas (type of M. atropurpureus Murrill), compatible with our collections except for the distinctly marginate lamellae of these two later. In fact, Singer (1976) considered M. sanguineus and M. atropurpureus synonyms under M. haematocephalus. However, would the ventricose (broad), distinctly marginate lamellae (edge concolorous with the pileus) be compatible with the protologue of M. haematocephalus? In Singer (1976), the lamellar edges are not concolorous with the pileus in mature basidiomata except “ sometimes ” near the pileus margin. It is possible that Singer’s concept was broadened to include the synonyms. Marasmius haematocephalus in Desjardin (1989), of collections from North America forests in the United States (Florida, North Carolina and Tennessee) along with collections from British Honduras, Cuba and Guyana, differs from the examined material only in having more numerous (10 - 15), broad (up to 2.5 mm), and marginate (pinkish red, deep reddish or violet red) lamellae (sometimes non-marginate) and a wider spore range (16 - 22 × [3.6 -] 4 - 5.6 µm). If the broad, marginate lamellae is a strong diagnostic characteristic in separating species, then there are M. haematocephalus with non-marginate lamellae vs M. sanguineus (synonym: M. atropurpureus) with marginate lamellae. Thus, the collections with marginate lamellae from North America examined in Desjardin (1989) may be M. sanguineus (or M. atropurpureus) instead of M. haematocephalus. Pegler (1983) should have had a broad (mix) concept on examined collections from Martinique, Trinidad, Dominique (including the type of M. sanguineus) and Sri Lanka (type of M. semipellucidus). The spore range was shorter (16 - 20 × 3 - 4.5 µm) than those of collections from São Paulo (18 - 22 × 3.7 - 4.5 µm) in Pegler (1997), this later more compatible (the purplish red pileus) with our collection as well as in Dennis (1970) from Venezuela (bright purple pileus with 19 - 21 × 3 - 4 µm basidiospores). With the finetuned concept of M. haematocephalus (s. str.) in this study, its various heterotypic synonyms listed in Singer (1976) and other need to be carefully reevaluated in future studies. By revising their protologues (Appendix), none of them seems morphologically compatible with M. haematocephalus s. str. except for the illegitimate M. vinosus Beeli from Africa. In Brazil, according to the speciesLink – Virtual Herbarium of the Centro de Referência em Informação Ambiental, CRIA (splink. org. br, accessed in November 17 th, 2019), 241 collections determined as M. haematocephalus are currently deposited in 23 Brazilian herbaria, collected from the states of Amazonas, Bahia, Espírito Santo, Maranhão, Minas Gerais, Pará, Paraíba, Pernambuco, Piauí, Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul, Rondônia, Roraima, Santa Catarina and São Paulo (Brazil). A review of these material is due in another paper considering the M. haematocephalus (s. str.) or M. rubicundus system. Singer (1976) recognized nine varieties in M. haematocephalus based on subtle distinctions and overlapping morphological characteristics: 1) atroviolaceus Singer; 2) haematocephalus; 3) leucophyllus Singer; 4) macrocephalus Singer; 5) oenechinus Singer; 6) pseudotageticolor Singer (partly marginate lamellae, red); 7) purpureomarginatus Singer (marginate lamellae, garnet red); 8) rubicundus Singer; and 9) transiens Singer. All these varieties are from the Neotropical realm, mostly from forests of South America. Except for the 8 th, these taxa need to be reevaluated with fresh collections. The sixth and seventh may be morphologically compatible with those in Desjardin (1989) with marginate lamellae. In Pegler (1987), the supposed (seems not formalized) additional M. haematocephalus var. obscurior Berk. & Curtis from Cuba has pileus browner than the typical form and with marginate lamellae, more compatible with M. hypophaeus. Desjardin (1991) proposed M. haematocephalus var. anomaloides Desjardin, of collections from Tennessee, United States. According to the current broad morphological concept, M. haematocephalus has been reported from various localities in South and Central America including some Caribbean islands (Neotropics), and less frequently from southern continental and insular North America (Nearctic) (Singer 1965, 1976; Dennis 1970; Pegler 1983, 1997; Desjardin 1989), Tropical Africa including Madagascar (Singer 1964; Pegler 1977; Antonín 2007; Shay et al. 2017), Tropical Asia (Petch 1948; Pegler 1986; Desjardin et al. 2000; Tan et al. 2009; Wannathes et al. 2009) and Oceania (Desjardin & Horak 1997). These collections need to be reevaluated given these new data and analyses, and a better understanding of the species complex. Marasmius haematocephalus may be confused with M. asiaticus, M. pallescens, M. panerythrus Singer, M. pulcherripes, and M. rhodopurpureus Antonín, R. Ryoo & H. D. Shin. Marasmius asiaticus differs by having “ less violet ” pileus, less numerous lamellae (7 - 8), slightly larger basidiospores (18.5 - 23 [- 25] × 3.5 - 5 [- 6] µm) and a mottled pileipellis (Tan et al. 2009; as M. distantifolius). Maramius pallescens differs by having a pale red pileus, shorter basidiospores (11 - 17 µm in length) and mottled pileus (Singer 1976). Marasmius panerythus and M. pulcherripes diverges by having shorter basidiospores (13 - 14 µm in length [Singer 1976] and [11 -] 12 - 15 [- 16] µm [Desjardin 1989; Antonín et al. 2012], respectively); and M. rhodopurpureus also has shorter basidiospores (12 - 15 [- 16] µm) and shorter (35 - 42 µm), non-refractive, clavate pleurocystidia (Antonín et al. 2012). In the phylogenetics trees (Figs 1; 3), various strains named after M. haematocephalus in previous studies forming a pantropical distribution branched in multiple lineages (para- or polyphyletic), indicating them to be not a single but close, cryptic species in haemat _ cp 1. Based on combined nrLSU + nrITS data, M. haematocephalus s. str. (JO 533) branched in a distinct phylogenetic position as sister to M. auranticapitatus J. S. Oliveira, sp. nov. + M. rubicundus (Singer) J. S. Oliveira, stat. nov. (Fig. 4).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301E7127FF0B7F9CE0CCE295.taxon	description	(Figs 15 A; 16; 17) Differs from M. pallescens in having longer basidiospores (12 - 20 µm vs 11 - 17 µm) and pleurocystidia (37.5 - 75.5 µm vs 32 - 43 µm); from M. pulcherripes also in the longer basidiospores (12 - 20 µm vs 11 - 15 µm), shorter setulae on the pileipellis broom cells (up to 4 µm vs up to 10 µm) and is devoid of the peculiar red stipe apex and inner extremities of the lamellae; from M. rubicundus (Singer) J. S. Oliveira, stat. nov. mainly in having smaller basidiospores, 12 - 20 × 3 - 4 µm (xm = 16.3 [± 2.4] × 3.6 [± 0.3] µm) vs (15.4 -) 16.2 - 21.7 (- 22) × 2.7 - 4.7 (- 5) µm (xmm = 19 [± 0.5] × 3.8 [± 0.2] µm); differs from all in the peculiar dimorphic Siccus-type broom cells in the pileipellis: 1) thick-walled, with pale chestnut lilac lumen, setulae apical, short, often coarse; 2) in transition to Rotalis-type broom cells, thin-walled.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301E7127FF0B7F9CE0CCE295.taxon	materials_examined	HOLOTYPE. — Brazil. São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, 3. III. 2011, J. J. S. Oliveira & F. Karstedt JO 352 (holo-, SP [SP 445510]!).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301E7127FF0B7F9CE0CCE295.taxon	etymology	ETYMOLOGY. — Based on the rose pileus. MYCOBANK. — MB 842538.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301E7127FF0B7F9CE0CCE295.taxon	description	HABIT AND SUBSTRATE. — Marasmioid (Figs 15 B; 16 A), gregarious, on dead eudicotyledonous sticks in the forest litter. DESCRIPTION Pileus (Figs 15 B; 16 A) 2 - 21 mm diam., convex, orbicular, sulcate to deeply sulcate, center flat, somewhat wrinkled when mature, margin decurved to plane, edge entire to slightly crenate; grayish pink to lilac (N 40 Y 10 M 60), becoming pale pink or rose (N 40 Y 20 M 50), center purple or dark pink (N 90 Y 00 M 80); membranous, context thin (<1 mm); glabrous, dry, dull to semi translucent, subvelutinous, non-hygrophanous. Lamellae (Figs 15 B; 16 A) Free to adnate, distant, L = 12, equal, simple, l = 0 (- 1), opaque, smooth, whitish pink (N 00 Y 20 M 20), edge even, non-marginate, interlamellar hymenium concolorous with the pileus. Stipe (Figs 15 B; 16 A) 8 - 40 × 0.3 - 0.7 mm, central, filiform, thin, equal, with circular caliber, chitinous, hollow; apex concolorous with the lamellae, becoming dark brown to almost black elsewhere, glabrous, smooth, with a silky bright; with a scarce, cream, tomentose basal mycelium. Odor Not distinctive. Basidiospores (Fig. 16 B) 12 - 20 × 3 - 4 µm (xm = 16.3 [± 2.4] × 3.6 [± 0.3] µm, Qm = 4.5 [± 0.6], n / s = 32 / 1), oblong, widely variable in length, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia Not observed. Basidioles (Fig. 16 C) 17.5 - 26.3 × 5.4 - 8.8 µm, cylindrical to clavate, sometimes tapering toward the apex, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia (Fig. 16 D) 37.5 - 75.5 × 6.3 - 10.6 µm, abundant, conspicuously projecting, cylindrical, clavate, sometimes ventricose, generally with tapered apex, or capitate, some worm-like to irregular in outline, apex easily breakable, smooth, slightly fuscous, yellowish, thin-walled, refractive, inamyloid. Cheilocystidia (Fig. 16 E) In form of Siccus-type broom cells, some apparently in transition to Rotalis-type similar to the broom cells of the kind 2 in the pileipellis; main body 11.3 - 17.5 × 6.3 - 10 µm, clavate to pyriform, rarely cylindrical to ventricose, thin-walled, hyaline; setulae apical to little divergent to the laterals, erect, generally short, 0.6 - 3.8 × 0.5 - 1 µm, digitiform or verruciform, simple or branched, solid, pale brown, regular to somewhat irregular in outline, apex obtuse to slightly acute. Lamellar trama Dextrinoid, irregular, interwoven, hyphae cylindrical, 1.3 - 7.5 µm diam., regular or irregular in outline, branched, smooth, hyaline, thin-walled. Pileus trama Similar to the lamellar trama, hyphae 2 - 7 µm diam. Pileipellis Hymeniform, mottled, composed of Siccus-type broom cells, of two kinds: 1) abundant at the pileus center and abruptly less frequent from the mid disc to the marginal zone (Fig. 17 A); main body 5 - 17.5 × 7.5 - 11.3 µm, pyriform, turbinate to subclavate, sometimes apically flattened, thick-walled especially at the apex, with pale chestnut lilac lumen; setulae apical, erect, short, 1.3 - 4 × 0.8 - 1.3 µm, digitiform to verruciform, or vesiculose, sometimes branched, often coarse, regular to irregular in outline, solid, hyaline to pale brown, apex obtuse; and 2) more numerous at the mid disc and predominant at the marginal zone (Fig. 17 B), some apparently in transition to Rotalis-type broom cells, rarely almost devoid of apical diverticula; main body 8.8 - 20 × 5.6 - 12.5 µm, clavate, turbinate, ventricose to subpyriform, walls very thin, completely hyaline, difficult to be individualized; setulae apical and frequently divergent to the laterals, erect, generally short, 0.3 - 2.5 (- 4) × (0.2 -) 0.5 - 1 µm, vesiculose, verruciform to digitiform, solid, pale brown, apex obtuse and rounded, or irregular. Stipe trama Dextrinoid, especially in the internal hyphae, cortical hyphae parallel, cylindrical, regular in outline, 2.5 - 8.8 µm diam., wrinkled, dark brown, walls not clearly thick, or thin-walled; internal hyphae hyaline, 2.5 - 10 µm diam., thin-walled. Clamp connections Present in all tissues, except in the cortical hyphae of the stipe. REMARKS Marasmius roseus J. S. Oliveira, sp. nov. differs from M. haematocephalus mainly by having pale purplish pink to pale lilac pileus center, then pink or rose from the mid disc to the margin, by having widely variable length of basidiospores (12 - 20 µm), and by having a mottled pileipellis with two kinds of Siccus-type broom cells (one apparently in transition to Rotalis-type). It is 2.2 % and 0.5 % dissimilar to M. haematocephalus s. str. (JO 533) in the nrITS and nrLSU, respectively. Marasmius roseus is morphologically somewhat similar to M. pallescens sensu Singer (1976), especially in the light lilac pileus form (Dennis 1961). Singer (1976) also observed a mottled pileipellis in M. pallescens due to some broom cells with darker melleous apical setulae among paler broom cells. However, M. pallescens has shorter basidiospores (11 - 17 µm) and smaller pleurocystidia (32 - 43 × 10 - 15.2 µm). Marasmius roseus J. S. Oliveira, sp. nov. is also very close to M. rubicundus (Singer) J. S. Oliveira, stat. nov. (next taxon) in the pileus pigmentation, but this later varies more widely from pale red to rose, ruby pink, even purplish pileus while has basidiospores less variable in length and larger in average (xmm = 18.8 [± 1.3] × 3.8 [± 0.2] µm), and non-mottled pileipellis composed of regular Siccus-type broom cells. Marasmius pulcherripes is similar especially by having similar pileus pigmentation and mottled pileipellis due to thickwalled, purplish broom cells among more regular ones in the pileipellis (Gilliam 1976; Antonín et al. 2012). However, it differs from M. roseus J. S. Oliveira, sp. nov. mainly by having shorter basidiospores (11 - 15 µm) and longer apical setulae (up to 10 µm) on the broom cells of the pileipellis beside the typical stipe pigmentation. None of the heterotypic synonyms under M. haematocephalus listed in Singer (1976) nor their varieties seem conspecific with M. roseus J. S. Oliveira, sp. nov. Marasmius haematocephalus var. transiens Singer also has basidiospores largely variable in size ([12.5 -] 13 - 22 × 3.5 - 4.5 µm), but the pileus (3 - 12 mm diam.) color is completely different with no similar cells in the pileipellis (Singer 1965, 1976).	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301C711BFF787A4CE067E144.taxon	description	(Figs 15 B; 18)	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301C711BFF787A4CE067E144.taxon	description	ADDITIONAL EXAMINED MATERIAL. — Brazil. São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 16. XII. 2009, M. Capelari & L. A. S. Ramos 4567 (SP [SP 445931]!); M. Capelari & L. A. S. Ramos 4570 (SP [SP 446073]!); 14. I. 2010, J. J. S. Oliveira & M. Capelari JO 1 (SP [SP 446075]!); 16. III. 2010, J. J. S. Oliveira JO 43 (SP [SP 446069]!); 15. X. 2010, J. J. S. Oliveira & C. L. A. Pires JO 223 (SP [SP 446074]!); 16. X. 2010, J. J. S. Oliveira & C. L. A. Pires JO 228 (SP [SP 445448]!); J. J. S. Oliveira & C. L. A. Pires JO 230 (SP [SP 446076]!); 17. X. 2010, J. J. S. Oliveira & C. L. A. Pires JO 245 (SP [SP 446046]!); J. J. S. Oliveira & C. L. A. Pires JO 246 (SP [SP 445454]!); 5. XI. 2010, J. J. S. Oliveira JO 261 (SP [SP 446051]!); 7. XI. 2010, J. J. S. Oliveira & A. V. Costa JO 275 (SP [SP 446059]!); 7. XII. 2010, J. J. S. Oliveira, P. O. Ventura & A. V. Costa JO 281 (SP [SP 445464]!); J. J. S. Oliveira, P. O. Ventura & A. V. Costa JO 283 (SP [SP 446061]!); J. J. S. Oliveira, P. O. Ventura & A. V. Costa JO 295 (SP [SP 446063]!); J. J. S. Oliveira, P. O. Ventura & A. V. Costa JO 296 (SP [SP 445476]!); 9. XII. 2010, J. J. S. Oliveira, P. O Ventura & A. V. Costa JO 316 (SP [SP 445488]!); J. J. S. Oliveira, P. O. Ventura & A. V. Costa JO 318 (SP [SP 446048]!); J. J. S. Oliveira, P. O. Ventura & A. V. Costa JO 319 (SP [SP 446047]!); São Paulo City, Parque Estadual das Fontes do Ipiranga, 1. III. 2011, J. J. S. Oliveira & F. Karstedt JO 330 (SP [SP 446065]!); J. J. S. Oliveira & F. Karstedt JO 335 (SP [SP 446067]!); J. J. S. Oliveira & F. Karstedt JO 338 (SP [SP 446066]!); 18. X. 2011, J. J. S. Oliveira & P. O. Ventura JO 380 (SP [SP 446064]!); São Paulo City, Parque Estadual da Cantareira, Núcleo Engordador, 19. XII. 2011, J. J. S. Oliveira & M. Capelari JO 445 (SP [SP 445537]!); J. J. S. Oliveira & M. Capelari JO 446 (SP [SP 446054]!); J. J. S. Oliveira & M. Capelari JO 447 (SP [SP 446053]!); 9. II. 2012, J. J. S. Oliveira & M. Capelari JO 481 (SP [SP 446043]!); J. J. S. Oliveira & M. Capelari JO 482 (SP [SP 446055]!); J. J. S. Oliveira & M. Capelari JO 483 (SP [SP 445560]!); 16. II. 2012, J. J. S. Oliveira & M. Capelari JO 492 (SP [SP 446045]!); J. J. S. Oliveira & M. Capelari JO 514 (SP [SP 446071]!); J. J. S. Oliveira & M. Capelari JO 516 (SP [SP 446057]!); Iporanga City, Parque Estadual Turístico do Alto Ribeira, Núcleo Santana, 29. II. 2012, J. J. S. Oliveira & D. E. Desjardin JO 529 (SP [SP 445577]!); D. E. Desjardin DED 8675 (SP [SP 445665]!); Lageado, 29. II. 2012, J. J. S. Oliveira & D. E. Desjardin JO 534 (SP [SP 446077]!). MYCOBANK. — MB 842539. HABIT AND SUBSTRATE. — Marasmioid (Figs 15 A; 18 A), solitary to gregarious, on dried eudicotyledonous leaves and sticks in the forest litter.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301C711BFF787A4CE067E144.taxon	distribution	DISTRIBUTION. — Originally from La Paz and Beni, Bolivia, as M. haematocephalus var. rubicundus (Singer 1965, 1976), it is now known from Southeastern Brazil.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B45301C711BFF787A4CE067E144.taxon	description	DESCRIPTION Pileus (Figs 15 A; 18 A) 2.5 - 20.5 mm diam., conical, hemispherical to convex, sulcate, center flat to umbonate, smooth or somewhat wrinkled, margin decurved to almost straight, edge entire to crenate; sometimes pale pinkish beige or salmon color (N 10 Y 20 M 40 to N 10 Y 30 M 50), often “ Canna ” pink or rose (N 20 Y 00 - 20 M 40 - 60), tending to very pale pinkish lilac (N 20 Y 20 M 60), then vibrant or ruby pink (N 40 Y 50 M 80), pinkish red (N 20 Y 50 M 70, N 20 Y 60 M 70 to N 20 Y 70 M 70), or pinkish brown “ Egyptian red ” (N 40 Y 50 - 60 M 60), sometimes pink with a slight hue of yellow (N 30 Y 50 M 50), or many times pale red or “ Carnelian ” (N 60 Y 40 - 60 M 90 - 99), light reddish brown (N 50 - 60 Y 90 - 99 M 90), with center darker red (N 70 Y 60 M 99) or pinkish brown (N 50 Y 70 M 70), or dark lilac (N 70 Y 00 M 99) or dark pink (N 70 Y 40 M 90); membranous, context thin (<1 mm); glabrous, dry, dull, papyraceus to subvelutinous, non-hygrophanous. Lamellae (Figs 15 A; 18 A) Free, adnexed, or sinuate to narrowly adnate, distant to subdistant, L = 7 - 12, equal, narrow to broad, straight, cultriform to ventricose, simple, l = 0 (- 1), dull, smooth, white to cream (N 00 Y 10 M 00), or whitish pink (N 00 Y 10 M 10 or N 00 Y 40 M 10 - 20), edges even, non-marginate, interlamellar hymenium concolorous with the lamellae faces or partly concolorous with the pileus. Stipe (Figs 15 A; 18 A) 8 - 63 × 0.3 - 0.8 mm, central, filiform, thin, equal, sometimes with broader base, with circular caliber, compressed when dried, chitinous, hollow; apex pale pink (N 00 Y 10 M 10 or N 30 Y 50 M 80) becoming bronze brown (N 40 Y 60 M 50 to N 80 Y 70 M 40) to dark brown (N 80 Y 99 M 30), or almost black at the base, glabrous, smooth, with a silky bright; with a scarce, white, tomentose basal mycelium. Odor Not distinctive. Basidiospores (Fig. 18 B) (15.4 -) 16.2 - 21.7 (- 22) × 2.7 - 4.7 (- 5) µm (xrm = 18 - 19.7 × 3.5 - 4 µm; xmm = 19 [± 0.5] × 3.8 [± 0.2] µm; Qrm = 4.6 - 5.5; Qmm = 5 [± 0.3]; n / s = 30 / 28), exceptional spore size of JO 316 with 18.3 - 23.9 × 3.1 - 4.8 µm (xm = 21.3 [± 1.3] × 3.9 [± 0.4] µm; Qm = 5.4 [± 0.6]; n / s = 30 / 1), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia (Fig. 18 C) 21.3 - 36.3 × 6 - 7.5 µm, clavate, smooth, hyaline to slightly fuscous, thin-walled, 4 - sterigmate, inamyloid. Basidioles (Fig. 18 D) 22 - 28.8 × 5.6 - 8.8 µm, clavate, sometimes as cystidioles, smooth, hyaline to slightly fuscous, inamyloid. Pleurocystidia (Fig. 18 E) (19.5 -) 25.6 - 71.7 × 5 - 12.3 µm, broadly clavate, capitate, almost mucronate or acuminate, with a conical apex, or ampullaceous, or occasionally with apical, serial, slight constrictions, ending with a capitule, smooth, slightly fuscous, thin-walled, refractive, inamyloid, abundant. Cheilocystidia (Fig. 18 F) Similar to the Siccus-type broom cells of the pileipellis; main body 11.3 - 20 × 5.6 - 10.6 µm, clavate to slightly turbinate, wall somewhat thick, hyaline; setulae more strictly apical, rarely somewhat divergent, erect, 2 - 7.5 × 0.5 - 1 µm, cylindrical, digitiform or needle-like, simple, solid, hyaline to pale brown, regular in outline, apex obtuse to slightly acute. Lamellar trama Dextrinoid, irregular, interwoven, hyphae cylindrical, 1.5 - 10 µm diam., regular in outline, branched, hyaline, thin-walled, smooth. Pileus trama Dextrinoid, similar to the lamellar trama, hyphae 2 - 10 µm diam. Pileipellis Hymeniform, composed of Siccus-type broom cells (Fig. 18 G), abundant, brown when grouped, bleaching in KOH solution; main body 7.5 - 20 × 6.3 - 11.3 µm, clavate, turbinate, sometimes branched, ventricose to irregular in outline, hyaline, thin-walled to slightly thick-walled, weakly dextrinoid; setulae apical, erect, 2 - 7.5 × 0.6 - 1 µm, cylindrical, digitiform or needle-like, simple, regular in outline, or slightly contorted, solid, hyaline to pale brown, apex obtuse to slightly acute. Stipe trama Dextrinoid especially the internal hyphae and those of the stipe apex, cortical hyphae parallel, cylindrical, regular in outline, 3.8 - 8 µm diam., smooth, dark chestnut brown, yellowish brown when separated, thick-walled; internal hyphae regular in outline, 2.5 - 6.5 µm diam., parallel. Clamp connections Present in all tissues, except in the cortical hyphae of the stipe. REMARKS The type collections were not examined in the present study as we had no reply from LIL herbarium in this pandemic time. However, Singer (1965, 1976) provided enough morphological data to unambiguously determine the examined specimens from São Paulo as authentic representatives of M. haematocephalus var. rubicundus. The pigmentation and size of the pileus, the number of lamellae, and the size of the basidiospores (15 - 21 × 4 - 4.8 µm in Singer [1976]) match the examined material. Marasmius rubicundus is 0.7 - 0.9 % (nrITS) and 0.5 - 0.7 % (nrLSU) dissimilar to M. haematocephalus, but cladogenesis reveals vicariance in sympatry. In the phylogenetic trees (Figs 1; 3) of nrITS, M. rubicundus (Singer) J. S. Oliveira, stat. nov., M. auranticapitatus J. S. Oliveira, sp. nov., and M. haematocephalus are embedded within haemat _ cp 2 a. In multilocus analyses, nrITS + nrLSU (Fig. 4) and nrITS + rpb 2 + ef 1 - α (Fig. 5), three species were resolved within this complex: M. rubicundus (Singer) J. S. Oliveira, stat. nov. sister to M. auranticapitatus J. S. Oliveira, sp. nov. with high support, and paraphyletic to M. haematocephalus. Morphologically, M. rubicundus (Singer) J. S. Oliveira, stat. nov. is very similar to M. haematocephalus, only more conspicuously distinct by having a lighter pileus pigmentation and slightly smaller basidiospores (x mm = 18.9 [± 0.5] × 3.8 [± 0.1] µm, Qmm = 5 [± 0.2] vs xmm = 20.5 [± 0.3] × 3.9 [± 0.2] µm, Qmm = 5.2 [± 0.1]). Marasmius haematocephalus s. str. regards the “ rubro - sanguineo ” of its protologue as deep or dark blood red or vinaceous (Singer 1958). In fact, the separation of M. rubicundus (Singer) J. S. Oliveira, stat. nov. and M. haematocephalus is very narrow in a broad sense, as they are nearly identical in other morphological characteristics. The species divergence is only evident in multilocus phylogenetic analyses where M. rubicundus (Singer) J. S. Oliveira, stat. nov. is rather closer to M. auranticapitatus J. S. Oliveira, sp. nov. (Figs 4; 5). The exceptional spore sizes of JO 316, possibly a hybrid, strenghtens the plausibility of the most recent common ancestor between them. Although speciation is demonstrated, this ancestry trace or possible partial intercompatibility and gene flow are elements of a recent divergence. The strain JO 316 seems in a transitional / intermediate position (Fig. 4) along with other strains of M. rubicundus (Singer) J. S. Oliveira, stat. nov., but the basidiospores are more compatible with M. auranticapitatus J. S. Oliveira, sp. nov .. JO 316 and JO 226 highlights the powerful phylogenetic signal of spore sizes, an evident correlation between this characteristic and the genotype behind the species concepts. The pale red pileus in M. rubicundus (Singer) J. S. Oliveira, stat. nov. is consistent with M. pallescens, but this later has shorter (11 - 17 µm) basidiospores (Singer 1976). For more color pictures showing variation of pileus pigmentation of M. rubicundus (Singer) J. S. Oliveira, stat. nov., see Appendix (Figs S 7 - S 11). It is not similar to any of the heterotypic synonyms of M. haematocephalus listed in Singer (1976) but M. rhodocephalus Fr. (Fries 1851; Appendix) with type presumed lost but connection is not sharp.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453020711CFC247CBFE09AE4A6.taxon	discussion	REMARKS	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453027711CFF32785DE3BEE186.taxon	description	Basidiomata marasmioid (umbrella-like), small- to medium-sized. Pileus membranous, campanulate to convex, smooth to shal- lowly sulcate, papillate (1.4 - 33 mm diam). Lamellae distant to subdistant. Stipe thin-cylindrical, dark-coloured, with a scarce basal mycelium. Basidiospores medium-sized, oblong (11 - 17 µm long, xm = 12.5 - 15.3 µm, Qm = 3.1 - 4.2). Pleurocystidia present (not clear in some cases), small- to medium-sized (up to 65 µm long). Pileipellis composed of Siccus-type broom cells. Habit solitary to gregarious, on dead, eudicotyledonous leaves and twigs.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453027711CFF32785DE3BEE186.taxon	materials_examined	TYPE SPECIES. — Marasmius pulcherripes.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453027711CFF32785DE3BEE186.taxon	discussion	REMARKS Marasmius pulcherripes was originally described from United States. Some strains determined after this species from Northwest Arkansas (MT 032485) and Boston Harbor Islands (MF 161270; Haelewaters et al. 2018), United States, branched in clade Pulcherripes (Fig. 1). This placement indicates the plausibility of the series.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453027711DFEF27B7DE1CFE2A4.taxon	distribution	DISTRIBUTION. — Originally described from Puerto Rico (Murrill 1915), it is also reported from Venezuela (Dennis 1961; Singer 1965, 1976; Pegler 1983) and Martinique (Pegler 1983). If the identity of this “ ruby pink form ” is confirmed as conspecific, this would be the first record from Brazil.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
9A684B453027711DFEF27B7DE1CFE2A4.taxon	description	DESCRIPTION Pileus (Figs 15 C; 19 A 1, A 2) 5 - 15 (- 20) mm diam., obtusely conical to broadly convex, or plano-convex, sulcate, center flat or slightly umbonate, shallowly depressed when fully matured, margin decurved to applanate, edge entire to slightly crenate; at first ruby overall (12 E 7 - 8), then remaining ruby pink (N 30 Y 30 - 50 M 99) to pinkish red (N 30 Y M 90), or fainting to greyish ruby (12 D 6 - 7), with a dark purplish red (N 70 Y 40 - 60 M 99) or ruby (12 E 7 - 8) center; membranous, context thin (<1 mm); glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae (Figs 15 C; 19 A 1, A 2) Free to adnexed, few toothed seceding, distant, L = 6 - 16, equal, broad (up to 1.5 mm), slightly ventricose, simple, l = 0 (- 1), opaque, smooth, white, edges even, non-marginate, interlamellar hymenium paller than the pileus or ruby. Stipe (Figs 15 C; 19 A 1) 19 - 55 × 0.3 - 0.8 mm, central, thick filiform, equal, or broadening toward apex, with circular caliber, chitinous, hollow; apex whitish pink (N 10 Y 20 - 40 M 20), abruptly becoming reddish brown (N 80 Y 99 M 60 - 80) to dark brown (N 90 Y 99 M 80 - 99), or almost black at the base, glabrous, smooth, with a silky bright; subinsititious or with a scarce, white, tomentose to strigose basal mycelium. Odor Not distinctive. Basidiospores (Fig. 19 B) (11.1 -) 12.3 - 17.3 × (3.2) 3.7 - 5 µm (xrm = 14.7 - 15 × 4.2 µm; xmm = 14.9 [± 0.2] × 4.2 µm; Qrm = 3.6; Qmm = 3.6; n / s = 30 / 2), oblong, clavate, subfusoid to fusoid, smooth, hyaline, thinwalled, inamyloid. Basidia (Fig. 19 D) 20 - 26.7 × 4.8 - 7.4 µm, clavate, smooth, hyaline, thin-walled, with four short, verruciform sterigmata, inamyloid. Basidioles (Fig. 19 C) (15.2 -) 18.7 - 28 × (3.8 -) 4.5 - 6.9 (- 7.5) µm, clavate, smooth, hyaline, inamyloid. Pleurocystidia (Fig. 19 E) 31.4 - 47 × 7.2 - 10.6 µm, mostly clavate, or broadly clavate, some slightly lageniform or ventricose, sometimes apically capitate, papillate or mucronate, or with digitifom projection, occasionally wavy with shallow constrictions, smooth, hyaline to somewhat fuscous, thin-walled, refractive, inamyloid. Cheilocystidia (Fig. 19 F) Similar to the Siccus-type broom cells of the pileipellis, but hyaline and with thinner walls; main body 7.6 - 19.8 × 5.2 - 9.9 µm, clavate to slightly turbinate, seldom branched or lobulate, thin-walled, inamyloid; setulae apical, erect, 3.5 - 7.1 × 0.5 - 1.1 µm, cylindrical or filiform, needle-like, regular in outline, simple, solid, hyaline, apex acute. Lamellar trama Strongly dextrinoid, irregular, interwoven, hyphae cylindrical, 1.2 - 6.1 µm diam., regular in outline, hyaline, smooth, thin-walled. Pileus trama Dextrinoid, similar to the lamellar trama, hyphae 2.8 - 6.9 (- 8.7) µm diam., branched. Pileipellis Hymeniform, composed of Siccus-type broom cells (Fig. 19 G), pale brown, but with rare more deeply pigmented cells, bleaching in KOH solution; main body 9 - 18.6 × 4.1 - 9.5 (- 15) µm, cylindrical thin, clavate, turbinate, sometimes almost pedicellate, pale brown, later hyaline, thin-walled to slightly thick-walled (firm-walled), inamyloid or weakly dextrinoid; setulae apical, erect, 2.5 - 7.7 × 0.7 - 1.5 µm, cylindrical or filiform, needlelike, rarely digitiform, simple, regular in outline, solid, pale brown, apex tapered, obtuse to mostly acute. Stipe trama Dextrinoid, some strands apparently inamyloid, stipitipellis and cortical hyphae parallel, packed, cylindrical, regular in outline, 2.6 - 9 µm diam., sometimes branched, smooth, those of the superficial layer highly melanized, dark chestnut brown, brown at the cortex, thick-walled; internal hyphae regular in outline, 2.6 - 6.8 µm diam., hyaline, thin-walled. Clamp connections Present in all tissues. REMARKS Murrill (1915) would have named Marasmius pallescens based on the pale-red pileus (5 - 8 mm broad), fading to isabelline on drying. Dennis (1961) reported the species from Venezuela with lilac pileus (7 mm diam.). Singer (1965, 1976) studied both the type from Puerto Rico (18 August 1912, J. R. Johnston 556) and the collection from Venezuela (20 June 1958, Dennis 117) and reported a pale red or light lilac (5 - 8 mm broad.) pileus when fresh (Murrill 1915; Dennis 1961), but reddish brown when dried. Pegler (1983) examined the same collections, adding a third from Dominique, and reported a “ Pale Flesh Colour ” pileus (5 - 15 mm diam.), sometimes paling to buff but retaining a slightly darker disc. All of them agreed for the campanulate sulcate pileus and distant, white, about 10 lamellae (10 - 14 in Pegler 1983) in the protologue. These collections from Brazil agree in nearly all characteristics but differs in the ruby pink to pinkish red (Fig. 15 C), larger pileus (11 - 15 mm diam.). The size of the basidiospores perfectly agree with Singer (1976) and Pegler (1983), but those in Dennis (1961) are shorter (11 - 14 × 3.5 µm); the shape and size of pleurocystidia are nicely compatible. The size of basidia and basidioles is slightly larger in our collections. Despite the distinctions mentioned above, the examined collections match the morphological concept of the species. Singer (1976) commented that “ at first sight one might be inclined to consider this species one of the numerous color variants of Marasmius haematocephalus ”. Indeed, M. pallescens seems very similar to M. haematocephalus, especially considering these possible collections from Brazil. However, both basidiospores and pleurocystidia are clearly shorter and the lamellae are broader and ventricose in M. pallescens. This species is also similar to M. panerythrus Singer, but this later differs by having more deeply pigmented pileus (purple red with paler “ copper leaf ” margin), by having marginate, pinkish lamellae, by having smaller basidiospores (13 - 14.7 × 3.5 - 4.2 µm), and by having larger pleurocystidia (20 - 62 × 6.8 - 9.8 µm) (Singer 1976). Marasmius cf. pallescens is similar to M. pulcherripes “ the pinkish-red form ” (Desjardin 1989), and originally described from Nearctic (New York, United States). Marasmius pulcherripes, however, has more numerous lamellae (15 - 16), more colorful stipe, slightly smaller basidiospores ([10.4 -] 12 - 16 × 3.2 - 4.6 µm), and regular lamellar trama. Antonín et al. (2012) reported M. pulcherripes from South Korea with marginate lamellae and larger pleurocystidia (37 - 65 × 7.0 - 12 µm), and this branched as a distinct lineage closer to M. siccus species complex (siccus _ cp 1) than to M. haematocephalus species complex (haemat _ cp 1) in Fig. 1. Unfortunately, no DNA sequences were obtained from the examined collections here as M. cf. pallescens. Because of the morphological similarity, one may predict it would branch closely related to M. pulcherripes sensu Antonín et al. (2012). Based on morphology, M. cf. pallescens is classified in ser. Pulcherripes (Oliveira et al. 2020) along with M. pallescens, M. panerythrus and M. rhodopurpureus.	en	S, Jadson José, Oliveira, ouza de, Capelari, Marina, Margaritescu, Simona, Moncalvo, Jean-Marc (2022): Disentangling cryptic species in the Marasmius haematocephalus (Mont.) Fr. and M. siccus (Schwein.) Fr. species complexes (Agaricales, Basidiomycota). Cryptogamie, Mycologie 20 (5): 91-137, DOI: 10.5252/cryptogamie-mycologie2022v43a5, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a5
