Marasmius haematocephalus Fr.

Marasmius haematocephalus (Mont.) Fr. (Figs 13; 14) Epicrisis Systematis Mycologici, seu Synopsis Hymenomycetum: 382 (Fries 1838). — Type: Brazil. Rio de Janeiro State, Rio de Janeiro City, Auguste de Saint Hilaire, holotype not found (Singer 1976), or none located (Desjardin 1989), or not extant (Robert et al. 2013), or supposedly found in the PC herbarium (V. Antonín, pers. comm.). Agaricus haematocephalus Mont., Annales des Sciences naturelles, Botanique, séries 2, 8: 369 (Montagne 1837). Androsaceus haematocephalus (Mont.) Pat., Journal de Botanique (Morot) 3 (20): 336 (Patouillard 1889). Chamaeceras haematocephalus (Mont.) Kuntze, Revisio generum plantarum (Leipzig) 3 (3): 456 (Kuntze 1898). 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 JO533 (epi-, designated here, SP[SP 445580]!), nrITS (ON502673) and nrLSU (ON502729). 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 JO507 (SP[SP 446044]!); Iporanga City, Parque Estadual Turístico do Alto Ribeira, Núcleo Santana, 04.XII.2011, J.J.S. Oliveira JO435 (SP[SP 446079]!); 29.II.2012, J.J.S. Oliveira & D.E. Desjardin JO527 (SP[SP 446068]!); Amazonas State, Novo Airão City, Urubuquara, Rio Unini, 24.X.2019, J.J.S. Oliveira & Francisco JO1363 (INPA[INPA289980]!). HABIT AND SUBSTRATE. — Marasmioid (Figs 13; 14A 1), close, on dried eudicotyledonous petioles in the forest litter. 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. DESCRIPTION Pileus (Figs 13; 14A 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; 14A 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; 14A 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. 14B) (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. 14C) 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. 14D) (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. 14E) 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. 14F) 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. 14G), 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. 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 × 4.4-5.6µm, 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 lamell

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