Areas sampled

Collections were obtained from four areas: 1) Reserva Biológica de Paranapiacaba (S23°46′00″–S23°47′10″ and W46°18′20″–W46°18′40″), a predominantly mountainous area, close to the coast, covered by Ombrophilous Dense Forest up to 891 m in altitude (Domingos et al. 2000, Xavier et al. 2008), with mean annual rainfall of 3 381 mm and mean temperature around 17.9 °C (Domingos et al. 2000); 2) Parque Estadual da Cantareira, Núcleo Engordador (S23°24′11.89″ and W46°35′12.29″), an area covered by Seasonal Semideciduous Forest of humid mesothermal climate up to 1 215 m in altitude (Xavier et al. 2008), mean annual rainfall 1 545 mm (Ventura et al. 1966) and temperatures 14.3–18.2 °C (Secretaria do Meio Ambiente 2000); 3) Parque Estadual das Fontes do Ipiranga (S23°38′08″–S23°40′18″ and W46°36′48″–W46°38′00″), an urban park of preserved Seasonal Semideciduous Forest (Pivello & Peccinini 2002) reaching 798 m in altitude on the Atlantic plateau (Xavier et al. 2008), mean annual rainfall 1 368 mm and mean annual temperature 23.9 °C (Santos & Funari 2002); 4) Parque Estadual Turístico do Alto Ribeira (S24°16′40″–S24°38′30″ and W48°27′20″–W48°44′00″) with a karst mountainous residual landscape (Godoy 2001) predominantly covered by Ombrophilous Forest (Ivanauskas et al. 2012), and a humid Cfb-Subtropical climate (Köppen 1948), with mean temperatures from 18–22 °C (Secretaria do Meio Ambiente 2000), and annual rainfall from 1 500–2 000 mm (Lepsch et al. 1990).

Morphological analyses and taxonomy

Key for the 24 taxa of Marasmius sect. Globulares sensu Antonín & Noordeloos (the Globulares-Sicci complex) from the Brazilian Atlantic Forest

• 1. Pileipellis composed strictly of smooth cells . . . . . . . . . . . 2

• 1. Pileipellis consisting mainly of Siccus-type broom cells . . 3

• 2. Pileus yellowish orange, becoming cream with a ochraceous disc, 45–81 mm diam, pleurocystidia 55–118 μm long . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. silvicola

• 2. Pileus pale brown, 7–37 mm diam, pleurocystidia 43–87 μm long . . . . . . . . . . . . . . . . . . . . . . . . . M. pallidibrunneus

• 3. Stipitipellis bearing cylindrical clavate to setoid caulocystidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

• 3. Stipitipellis without smooth, clavate to cylindrical caulocystidia, but broom cells may be present . . . . . . . . . . . . . . . . 6

• 4. Pileus orange, up to 11 mm diam, basidiospores 13.8–18 μm long, caulocystidia setoid, elongate, up to 415 μm long, pileipellis made up only by Siccus-type broom cells . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. atrorubens

• 4. Pileus mainly white to cream, up to 66 mm diam, basidiospores up to 10 μm long, caulocystidia shorter and non-setoid, pileipellis formed either by smooth cells or by Siccus-type broom cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

• 5. Pileus pale cream, cheilocystidia of two types, both smooth and broom cells, caulocystidia rare and short near the stipe base, up to 21.3 μm long, basidiospores up to 9 μm long .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. congregatus

• 5. Pileus mainly pure white, cheilocystidia hyphoid, caulocystidia numerous covering the whole stipe surface, up to 62 mm long, basidiospores up to 10.4(–11) μm long . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pseudoniveoaffinis

• 6. Pleurocystidia present . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

• 6. Pleurocystidia absent . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

• 7. Lamellae intervenose or veined . . . . . . . . . M. venatifolius

• 7. Lamellae simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

• 8. Pileus olivaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

• 8. Pileus not olivaceous, varying between yellow and orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

• 9. Pileus up to 23 mm diam, olivaceous-brown, basidiospores 6.3–10 μm long . . . . . . . . . . . . . . . . . . . . M. altoribeirensis

• 9. Pileus up to 12 mm diam, yellowish olive, basidiospores 9.3–12.5 μm long . . . . . . . . . . . . . . . . . . M. luteoolivaceus

• 10. Pileus 21–45 mm diam, brownish orange, basidiospores 7.5–11.3 μm long. . . . . . . . . . . . . . . . . . . M. spegazzinii

• 10. Pileus up to 20 mm diam, yellow to orange, basidiospores > 11 μm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

• 11. Pileus up to 12 mm diam, lamellae marginate, orange, basidiospores 12.5–17.5 μm long . . . . . . . . . . . M. anomalus

• 11. Pileus reaching 20 mm diam, lamellae non-marginate, basidiospores up to 14 μm long. . . . . . . . . . . . . . . . . . 12

• 12. Pileus yellow with a slight orangish tint, setulae of broom cells of the pileipellis no more than 7.5 μm long; more frequently on leaves . . . . . . . . . . . . . . . . M. suthepensis

• 12. Pileus bright orange, setulae of broom cells of the pileipellis reaching 14.3(–17.5) μm long; often on wood . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. neotropicalis

• 13. Lamellae strongly reticulate to merulioid-anastomosed . . . . . . . . . . . . . . . . . . . . . M. cladophyllus var. glaberripes

• 13. Lamellae non-reticulate or merulioid-anatomosed . . . 14

• 14. Pileus cream-coloured . . . . . . . . . . . . . . . . . . . . . . M. bellus

• 14. Pileus strongly pigmented . . . . . . . . . . . . . . . . . . . . . . 15

• 15. Pileus olivaceous grey, basidiospores 10–13.3 μm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. trinitatis

• 15. Pileus not olivaceous, basidiospores with different dimensions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

• 16. Basidiospores > 11 μm long . . . . . . . . . . . . . . . . . . . . . 19

• 16. Basidiospores < 11 μm long . . . . . . . . . . . . . . . . . . . . . 17

• 17. Pileus bright orange to dark reddish orange (fulvous), stipe trama with dimorphic hyphae (few vermiform conspicuous hyphae in a matrix of regular hyphae) . . . . . . . . . M. ruber

• 17. Pileus yellowish orange or pale orange (not fulvous), stipe trama without dimorphic hyphae (only regular hyphae) 18

• 18. Pileus trama heteromorphic, hyphae thin- to thick-walled, pileipellis not mottled, broom cells with regular apical setulae, 2–6.3 μm long . . . . . . . . . . . . . . . . . . . . M. leoninus

• 18. Pileus trama not heteromorphic, hyphae only thin-walled, pileipellis mottled, some broom cells with thick walls and very elongate setulae . . . . . . . . . . . . . . . . M. corrugatus

• 19. Pileus < 10 mm diam, pileipellis not mottled . . . . . . . . . 20

• 19. Pileus > 10 mm diam, pileipellis mottled . . . . . . . . . 22

• 20. Pileus up to 7.5 mm diam, stipe 33–60 mm long, basidiospores 10–14.4 μm long . . . . . . . . . . . . . . M. graminicola

• 20. Pileus up to 4 mm diam, stipe shorter, up to 15 mm long, basidiospores longer . . . . . . . . . . . . . . . . . . . . . . . . . . 21

• 21. Pileus 1.8–4 mm diam, stipe 13–15 mm long, basidiospores 14–19 × 5.4–8 μm . . . . . . . . . . . . . . . . M. hobbitii

• 21. Pileus 1–3 mm diam, stipe 1–2.5 mm long, basidiospores 12.3–17 × 3.8–5 μm . . . . . . . . . . . . . . M. ambicellularis

• 22. Pileus beige to pale brown, basidiospores 12.8–16 × 3.5–4 μm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. dimorphus

• 22. Pileus dark orange, basidiospores longer . . . . . . . . . . 23

• 23. Basidiospores 20–25.4 μm long . . . M. rhabarbarinoides

• 23. Basidiospores (12–)14.4–21.5 μm long . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. rhabarbarinus

1 — Marasmius sect. Globulares Kühner emend. Antonín & Noordel. group Globulares

[Sect. Globulares sensu Singer (1986)]

Marasmius pallidibrunneus J.S. Oliveira, sp. nov. — MycoBank MB811130; Fig. 3, ​,44

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Fig. 3

Marasmius pallidibrunneus (M. Capelari 4607). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. pleurocystidia; f. cheilocystidia. — Scale bars: a = 21 mm, b–f = 10 μm.

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Fig. 4

Marasmius pallidibrunneus (M. Capelari 4607). a. Globulares-type smooth cells of the pileipellis; b. stipitipellis. — Scale bars: 10 μm.

Holotype. Brazil, São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, on rotten debris of dicotyledonous tree, 3 Apr. 2013, M. Capelari 4706 (SP 445668!).

Etymology. Referring to the pileus with pale brown pigmentation.

Pileus 7–37 mm diam, shallowly sulcate, pale brown or buff brown. Lamellae adnate to adnexed, subdistant (18–21). Stipe 18–50 × 2–4 mm, cylindrical, entirely white, except dark brown at the base and with a white basal pad. Basidiospores 6.3–9 × 3–4.4 μm, lacrimoid to clavate. Pleurocystidia 43–87.5 × 6.3–11.3 μm, cylindrical, clavate, ampullaceous, ventricose to vermiform. Cheilocystidia 34–65 × 8–15 μm, ampullaceous to often bulboid-pedicellate, yellow to pale ochraceous. Pileipellis hymeniform, composed of dextrinoid Globulares-type smooth cells, 16.3–43.8 × 7.8–21.3 μm.

Pileus 7–37 mm diam, hemispheric to convex, sometimes tending to plane, shallowly to deeply sulcate, centre flat, margin incurved, decurved to straight, edge regular; whitish brown to pale brown (N₄₀Y₃₀M₂₀), or light brown (N_30–40Y₃₀M₂₀), darkening to ochraceous brown (N₆₀Y₅₀M₄₀) or dark rust brown (N₉₀Y₈₀M₅₀) with humidity change, with a central, rounded spot, which is ochraceous brown (N₆₀Y₅₀M₄₀) to dark rust brown (N₉₀Y₈₀M₅₀); thin-fleshy or thin-cartilaginous to almost membranous at the margin, context white (≤ 1.5 mm); surface glabrous, dry, dull, papyraceous, darkening after collected, non- or partly hygrophanous. Lamellae adnate to adnexed, subdistant, ventricose (4 mm broad), L = 18–21, equal, l = 1–2, opaque, smooth, cream (N₁₀Y₂₀M₁₀) to greyish beige (N₂₀Y₃₀M₂₀), or pale brown (N₃₀Y₃₀M₂₀), edge even, concolorous. Stipe 18–50 × 2–4 mm, central, cylindrical, slightly compressed at the middle, equal, cartilaginous, hollow; white or greyish white all over because of a fine mycelial vestiture on the surface, changing to brownish grey (near N₃₀Y₂₀M₁₀) when mature, and dark brown (N₈₀Y₉₉M₅₀) to fuliginous brown (N₈₀Y₂₀M₂₀) at the base; surface with mycelial covering or a fine tomentum, stamped with finger print, but with thin pubescence at the stipe base, sometimes fibrillose; with a white, cottony basal mycelial pad, and forming a subiculum over the substrate.

Basidiospores 6.3–9 × 3–4.4 μm (x_m = 7.9 (± 0.8) × 3.9 (± 0.3) μm; Q_m = 2.1 (± 0.2); n = 30), oblong, tear-shaped to short clavate, hyaline, smooth, thin-walled, inamyloid. Basidia 27–36.3 × 6.3–7.5 μm, cylindrical to clavate, smooth, hyaline, 4 sterigmata, thin-walled, inamyloid. Basidioles 27.5–36.3 × 4.4–6.3 μm, cylindrical to clavate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia 43–87.5 × 6.3–11.3 μm, cylindrical, clavate, ampullaceous, ventricose to vermiform, smooth, hyaline, base a little deepened in the subhymenium, apex at the same height or a little more projected above the basidia or basidioles apex, solely conspicuous when individualized, thin-walled. Cheilocystidia 34–65 × 8–15 μm, abundant, conspicuous, erect to frequently repent, clavate, hyphoid, ampullaceous to frequently sphero-pedunculate, with the apical portion generally vesiculose or inflated, few hyaline, generally opaque, yellow to pale ochraceous, thin- or thick-walled, sometimes seeming solid likely due to contents. Lamellar trama dextrinoid, irregular, composed of strongly interwoven, cylindrical hyphae, 1.3–7.5 μm diam, regular in outline, branched, smooth, thin-walled, hyaline. Pileus trama dextrinoid, irregular, composed of hyphae similar to those of the lamellar trama, 3–15 μm diam, some inflated. Pileipellis hymeniform, composed of Globulares-type smooth cells, 16.3–43.8 × 7.8–21.3 μm, vesiculose, clavate to pyriform, pedicellate, some flattened at the apex, others tending to be lobed, sometimes septate, hyaline, dextrinoid, thin- to slightly thick-walled. Stipe trama strongly dextrinoid, cortical hyphae parallel, cylindrical, 3.8–12.5 μm diam, regular in outline, branched, smooth, pale brownish yellow, with walls moderately thick, septate; internal hyphae hyaline, thin-walled. Stipitipellis with a tomentum-like covering of loose hyphae on the cortical layer (the cause of the pale pigmentation of the stipe), with many basidiospores adhered to the hyphae of this vestiture, some of them germinating, tomentum formed by filamentous extensions of cylindrical hyphal endings, clavate to broadly clavate, or irregular in outline, septate, hyaline, thin-walled, inamyloid; caulocystidia (if present) in form of terminal hyphae growing from this layer, but definitely not obvious. Clamp connections present in almost all tissues, not seen at the base of the cheilocystidia or smooth cells of the pileipellis.

Habit & Substrate — Gymnopoid, gregarious, on rotten debris of dicotyledonous tree (leaves, branches and twigs), debris connected by a white to cream-coloured subiculum.

DNA barcode — GenBank KP635186 (ITS1-5.8S-ITS2) and KP635141 (LSU).

Notes — This species is similar to M. cibarius (Singer 1976), which also has pileus darkening to fuliginous brown, lamellae tending to be sordid grey, and the stipe is pale and slightly fibrillose. However, M. cibarius strongly differs in having smaller basidiospores (5.5–6.5 × 2.8–3.8 μm) and lacking pleurocystidia. Marasmius pallidibrunneus also resembles M. heliomyces (Singer 1976) in the colour and size of the pileus, the pale stipe and basidiospores dimensions (6–8.8 × 2.8–3.8 μm), but the latter species differs by the absence of pleurocystidia.

Some resemblances between M. pallidibrunneus and M. brunneospermus are the white pruinose stipe surface (at least sometimes in the latter), the presence of long pleurocystidia (50–95 μm long in the latter), the slightly pigmented cheilocystidia (pale melleous in the latter) of somewhat similar shape, the smooth cells of the pileipellis, size and shape of the basidiospores, and the lamellae tinted brown. However, both species are different according to the macroscopic features of the pileus (colour, texture and surface aspects) and in the number of lamellae (Takahashi 1999), and the lamellae are not intervenose in the new species. Moreover, the basidiospores or spore print of M. pallidibrunneus are not brownish. The presence of fusoid pleurocystidia and well-developed cheilocystidia also differentiate M. brunneospermus from M. pallidibrunneus.

In the Dataset 2.4 tree (Fig. 1e), M. pallidibrunneus was basal to the subclades /wynneae and /brunneospermus without support. On the other hand, in the Dataset 3 tree (Fig. 2), M. pallidibrunneus is sister to the branch bearing M. silvicola, M. macrocystidiosus, M. fusicystidiosus and M. brunneospermus, with significant support.

Marasmius silvicola Singer in Singer & Digilio, Lilloa 25: 199. 1952 — Fig. 5, ​,66

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Fig. 5

Marasmius silvicola (JO357). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. cheilocystidia; f. Globulares-type smooth cells of the pileipellis. — Scale bars: a = 17 mm, b–f = 10 μm.

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Fig. 6

Marasmius silvicola (JO357). Pleurocystidia. — Scale bars: 10 μm.

Pileus 45–81 mm diam, convex, tending to plane, orbicular, striate, corrugated or slightly sulcate, centre flat or with a shallow depression, sometimes slightly umbonate, margin incurved to straight, or tending to revolute, edge regular, crenate to lacerate; brownish yellow to orangish (N₃₀Y₉₀M₄₀), fading to yellow or beige (N₂₀Y₆₀M₃₀) to almost cream (N₁₀Y₆₀M₂₀) at the margin, centre ochraceous (N₁₀Y₉₉M₅₀) to dark yellowish brown (darker than N₄₀Y₉₉M₅₀); membranous to thin-fleshy, context moderately thick (± 1.5 mm); surface glabrous, semi-humid to moist, sometimes dry, dull at the central disc, usually pellucid at the margin, smooth to papyraceous, hygrophanous. Lamellae adnate, adnexed to sinuate, sometimes descending from the stipe, somewhat close to subdistant, L = 18–20, broad (17 mm), regular, some weakly intervenose, l = 3–5, opaque, smooth, greyish yellow to sordid cream (N₀₀Y₂₀M₀₀), edge even, sometimes eroded or irregularly toothed, concolorous with the lamellar faces (darkening after losing water or when completely dried) and with the hymenium between lamellae. Stipe 75–87 × 3–5 mm, central or somewhat eccentric, cylindrical, equal or with a flared apex, sometimes compressed, with scarce, white, tomentose basal mycelium, forming a subiculum; cartilaginous, hollow; apex and middle cream (N₀₀Y₁₀M₈₀) or becoming brownish (N₂₀Y₄₀M₃₀ to N₄₀Y₅₀M₄₀) when touched, and dark greyish brown (N₅₀Y₅₀M₃₀) downward to the base; surface glabrous or with very scanty, white pubescence, smooth, opaque, fibrillose. Odour not distinguished.

Basidiospores 5.6–9 × 3–4.8 μm (x_rm = 6.7–7.6 × 3.4–3.8 μm, x_mm = 6.9 (± 0.5) × 3.6 (± 0.2) μm, Q_rm = 2, n/s = 30, s = 3), obovoid, rarely ellipsoid, generally subellipsoid, lacrimoid or amygdaliform, smooth, hyaline, thin-walled, inamyloid. Basidia 27.5–38.8 × 5–6.3 μm, clavate, elongate, smooth, hyaline, thin-walled, with 2–4 sterigmata, inamyloid. Basidioles (17.5–)22.5–35 × 3.8–6.3 μm, cylindrical to clavate, short to elongate, slender, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia generally very elongate, 55–118 × 8.8–13.8 μm, clavate or frequently fusoid, tapering upward to an obtuse or acute apex, base elongate and very narrow, deepened in the subhymenium, some opaque (either when in 5 % KOH or in 5 % NH₄OH or NH₃), yellowish to very pale melleous, or hyaline, present either on the faces or at lamellae edges (repent at the edge), smooth, thin-walled, metachromatic, inamyloid. Cheilocystidia in form of smooth cells, subclavate, pyriform, sometimes bilobed, or pedicellate, (8.8–)12.5–27.5 × 5.6–8.8(–14.5) μm, hyaline, with thinner walls, inamyloid. Lamellar trama dextrinoid, irregular, tending to regular at the trama core, hyphae interwoven, cylindrical or inflated, 2.5–12.5 μm diam, broad, smooth, hyaline, thin-walled. Pileus trama dextrinoid, irregular, with a loose core, subcutis and subhymenium more compact, hyphae interwoven, 2.5–20 μm diam, cylindrical or inflated, broad, smooth, hyaline, thin-walled. Pileipellis hymeniform, composed of Globulares-type smooth cells, clavate to pyriform, sometimes slightly pedi-cellate, or with variable shape, sometimes like segments of hyphal endings, 16–33 × (8.8–)10.4–18.8 μm, smooth, hyaline when isolated, brownish yellow when grouped (pigmented walls), inamyloid, thin-walled. Stipe trama dextrinoid, especially the inner hyphae; cortical hyphae parallel, cylindrical, 2.5–17.5 μm diam, regular in outline, melleous or pale yellow, smooth, walls thin to moderately thick; internal hyphae similar to those of the cortex, but disorganised. Stipitipellis without differentiated cells. Clamp connections present in most tissues, mostly inconspicuous, not observed in the stipe trama or on the pileipellis elements.

Habit & Substrate — Gymnopoid, gregarious, scattered or solitary, on trunks or rotten vegetal humus of dicotyledonous trees.

Distribution — The species was originally described from Argentina (Singer & Digilio 1952), later collected from Rio Grande do Sul State in Brazil, Ecuador, Bolivia (Singer 1976), and more recently from Indonesia (Desjardin et al. 2000). This is the first record from São Paulo State in Brazil.

Specimens examined. Brazil, São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, 23 Mar. 2011, J.J.S. Oliveira & P.O. Ventura JO357 (SP 445514!), J.J.S. Oliveira & P.O. Ventura JO362 (SP 445517!), 4 May 2011, J.J.S. Oliveira & P.O. Ventura JO366 (SP 445519!).

Notes — The examined samples correspond to the species as described by Singer (1976), differing only by having slightly larger basidiospores, reaching 9 μm long, though the longer spores may have come from 2-spored basidia. The cheilocystidia ((8.8–)12.5–27.5 × 5.6–8.8(–14.5) μm) resemble the smooth cells of the pileipellis rather than the pleurocystidia, and this is consistent with the second pattern found by Singer (1976): “or else more inconspicuous, vesiculose hyaline, smooth, 11–28 × 6–10 μm”. As the pleurocystidia also occur at the lamellar edge, they may be confused with the cheilocystidia. No other significant differences are found. Desjardin et al. (2000) recorded this species as having very large basidiomata, with pileus ranging from 50–90(–200) mm diam and stipe with 80–140 × 5–10 mm.

Marasmius silvicola is also comparable to M. goossensiae, M. nigrodiscus, M. notandus, M. ochraceus, M. phlebodiscus and M. trogioides, differing from all by having much larger pleurocystidia. Furthermore, M. goossensiae has smaller basidiospores, reaching 7.5 μm long and pleurocystidia only up to 72 μm long (Singer 1964, Pegler 1977); M. notandus has dry, smooth, some weakly striate pileus, larger basidiospores (reaching 11 μm long), and completely unpigmented and shorter pleurocystidia, up to 70 μm (Corner 1996); M. ochraceus has completely smooth pileus, smaller basidiospores (5.5–7.2 × 2.5–4.2 μm) and pleurocystidia with strongly greenish yellow content (Pegler 1986); M. phlebodiscus has a pileus with a reticulated central disc by wrinkles in the form of a web, shorter basidiospores (up to 6.5 μm long) and cheilocystidia with tapered apex (Desjardin & Horak 1997); and M. trogioides has a smooth to rugulose-reticulate, or to rivulose-rugulose pileus, stipe with dilated base, longer basidiospores (9–11.5 μm), pleurocystidia reaching 70 μm long and very elongate cheilocystidia, up to 80 μm long (Corner 1996).

Based on the Dataset 2.4 tree (Fig. 1e), M. silvicola is sister to M. nigrodiscus with high support forming the subclade /silvicola. In the Dataset 3 tree (Fig. 2), M. silvicola has a stable phylogenetic position basal to the ‘M. brunneuspermus + M. fusicystidiosus + M. macrocystidiosus’ branch, three species that similarly have elongate and apically tapered pleurocystidia.

2 — Marasmius sect. Globulares Kühner emend. Antonín & Noordel. group Sicci Singer ser. Atrorubentes Desjardin & E. Horak

[Sect. Sicci sensu Singer (1986)]

Marasmius atrorubens (Berk.) Sacc., Syll. Fung. (Abellini) 5: 560. 1887 — Fig. 7

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Fig. 7

Marasmius atrorubens (JO489). a1, a2. Basidiomata; b. basidiospores; c. basidioles; d. cystidioles; e. cheilocystidia; f. Siccus-type broom cells of the pileipellis; g. setoid caulocystidia. — Scale bars: a1 = 15 mm, a2 = 7 mm, b–g = 10 μm.

≡ Agaricus atrorubens Berk., London J. Bot. 1(3): 138. 1842.

≡ Androsaceus atrorubens (Berk.) Pat., J. Bot., Paris 11: 338. 1897.

Pileus 1.4–3 mm diam when immature, reaching 2.6–11 mm diam, hemispheric to slightly conical when young, remaining conical, conical-campanulate or even convex to plano-convex when mature, smooth, striate or very slightly sulcate at the margin, centre shallowly umbonate, margin incurved to rarely straight, edge regular to slightly crenate; dark ferruginous or orangish brown (darker than N₆₀Y₉₉M₈₀) and centre with an orangish brown papilla (N₄₀Y₉₉M₇₀ to N₆₀Y₉₉M₇₀) when immature, becoming orange to bright orange (N₂₀Y₉₉M_50–60), some paler orange (N₃₀Y₉₉M₅₀) when mature at the margin, fulvous orange (N₄₀Y₉₉M_60–70) at the centre, usually becoming ochraceous brown (N₆₀Y₈₀M₅₀) when dried; membranous, context pale cream, thin (< 1 mm); surface glabrous, dry, dull, papyraceous to subvelutinous, non-hygrophanous. Lamellae sinuate, adnexed or subfree, subdistant to subclose, L = 12–22, equal or unequal, regular, l = 2–4(–5), opaque, smooth, pale cream (N₀₀Y₁₀M₀₀), edge even, finely concolorous with the pileus, hymenium between the lamellae partly concolorous with the pileus. Stipe 4.3–73 × 0.3–1 mm, central, cylindrical, equal; semichitinous to semicartilaginous, hollow; apex concolorous with the lamellae or pale orange (N₁₀Y₇₀M₃₀) to pale yellow (N₀₀Y₅₀M₁₀), becoming ochraceous brown (N₅₀Y₉₉M₅₀), orangish brown (N₄₀Y_80–90M₇₀), reaching somewhat reddish brown (N₇₀Y₉₉M₇₀) near the base; surface opaque, pubescent, with fuliginous brown pubescences; strigose basal mycelium long-filamentous, cream to orange, sometimes scarce.

Basidiospores 12.5–18 × 3–4.6 μm (x_rm= 14.5–15.9 × 3.5–3.6 μm; x_mm = 14.8 (± 0.7) × 3.6 (± 0.1) μm; Q_rm = 4–4.4; Q_mm = 4.3 (± 0.2); n/s = 30; s = 3), oblong, clavate to very slightly fusoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 17.5–28 × 5.8–7.5 μm, clavate, hyaline, thin-walled, inamyloid; cystidioles, 21.3–30 × 3.8–6.5 μm, fusoid to subfusoid, with apex generally tapered and acute, rarely almost mucronate, some with a broadened apical portion, projecting 8 μm beyond the basidia and basidioles, smooth, hyaline, thin-walled. Cheilocystidia concentrated at the lamellar edge, but also dispersed toward the middle of the lamellar face, in form of Siccus-type broom cells, abundant, ochraceous or pale brownish orange, similar to those of the pileipellis, main body 8.8–22.5 × 6.3–7.8(–10) μm, clavate to slightly turbinate, rarely truncate or ventricose, or branched, regular in outline, hyaline when isolated, walls relatively thin; setulae apical, erect, short to slightly elongate, 1.6–7.8 × 0.5–1 μm, simple, cylindrical to filiform, or setoid, regular in outline, hyaline to pale yellow, apex acute to somewhat obtuse. Lamellar trama strongly dextrinoid, irregular, composed of interwoven hyphae, cylindrical, 1.5–10 μm diam, regular in outline, branched, hyaline, thin-walled. Pileus trama strongly dextrinoid, irregular, interwoven hyphae, cylindrical, 2–10 μm diam, regular or sometimes irregular in outline. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, pale brownish orange when grouped, hyaline with apex and setulae hyaline to light yellow when isolated, main body 10–25 × 5–9.4 μm, clavate to turbinate, sometimes ventricose, regular in outline, hyaline, with walls somewhat thick; setulae apical, erect, 1.5–6.3 × 0.6–1 μm, cylindrical, digitiform to spinule-shaped, regular in outline, simple, solid, hyaline to light yellow, apex obtuse to more frequently acute. Stipe trama strongly dextrinoid, cortical hyphae parallel, cylindrical, 2.5–7.5 μm diam, regular in outline, smooth, ochraceous brown or pale hazel, thick-walled; internal hyphae hyaline, 2.5–9 μm diam, thin-walled. Caulocystidia (or Caulosetae) short to very elongate, 35–415 × 4.5–17.5 μm, setoid, cylindrical, tapering at the apex, sometimes forming an inflated spherical portion at the apex or in the middle of the cystidia, apex acute, not branched, pale brown to brown, thick-walled. Clamp connections present in almost all tissues, except for the cortical hyphae of the stipe.

Habit & Substrate — Marasmioid, scattered or gregarious, growing on dried dicotyledonous leaves on the forest litter.

Distribution — Originally described from Suriname, it has also been recorded from Guadalupe, Venezuela, Ecuador (Singer 1976), Tanzania, Uganda (Pegler 1977), Martinique, Trinidad (Pegler 1983), Puerto Rico (Lodge 2003, Cantrell et al. 2006) and Brazil (Pegler 1997).

Specimens examined. Brazil, São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, 1 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO337 (SP 445501!); Parque Estadual da Cantareira, Núcleo Engordador, 30 Jan. 2012, J.J.S. Oliveira & V. Motato-Vásquez JO455 (SP 445906!); ibid., 16 Feb. 2012, J.J.S. Oliveira & M. Capelari JO489 (SP 445563!); Iporanga, Parque Estadual Turístico do Alto Ribeira, Núcleo Santana, 29 Feb. 2012, J.J.S. Oliveira & D.E. Desjardin JO528 (SP 445576!); ibid., D.E. Desjardin DED8684 (SP 445663!).

Notes — Marasmius atrorubens is the type species of the ser. Atrorubentes according to the traditional view. Singer (1976) described M. atrorubens var. cystidifer based on specimens with abundant, ventricose to tapered ‘fusoid’ cystidioles, 22–27 μm long, with a conical or mucronate apex, slightly granular surface or covered with hyaline incrustations, all hyaline or having a distinct yellowish and opaque content, thick-walled and refractive. The specimens examined also have ventricose to tapered (‘fusoid apex’) cystidioles, 21.3–30 μm long, with a conical-subacute to mucronate apex, but differ by being hyaline, thin-walled, smooth, and non-refractive. These cystidioles are mostly concordant with the description of the species by Puccinelli & Capelari (2009). Sequencing more collections across this range of variation should clarify the relevance of these characteristics in the species.

Pegler (1977), based on material from Africa, mentioned the presence of ventricose and fusoid cystidioles (22–26 μm long). However, in Pegler (1983), cystidia or cystidioles were apparently absent in specimens from Martinique and Trinidad, but with only fusoid basidioles instead, and the basidiospores were much smaller (10–15 μm long), so this identification is dubious. Later, Pegler (1997) observed basidiospores ranging from 13–18 μm long in material from São Paulo and this agrees with the basidiospore dimension originally described for M. atrorubens.

Marasmius atrorubens var. dumontii (Singer 1976) differs from the other varieties by not bearing cystidia/cystidioles in the hymenium, but elongate, hair-like, thick-walled structures in the pileipellis. This latter characteristic points to a strong resemblance between this variety and M. trichotus, which has long pileosetae and shorter basidiospores, ranging from 11–14 μm (Corner 1996).

In the subtree of the Group 2 (Fig. 1c), M. atrorubens grouped with M. trichotus, M. longisetosus, M. iras, M. xestocephalus and M. nummularius, forming the subclade ‘atrorubens’. Marasmius atrorubens appears close to M. trichotus but, despite the considerable morphological similarity, the subtending branch is unsupported (Fig. 1c). Nevertheless, both branches of M. atrorubens and M. trichotus are strongly supported, indicating that the presence of pileosetae in M. trichotus represents a strong delimitation and that they represent distinct taxa.

Marasmius congregatus Mont., Ann. Sci. Nat., Bot., ser. 4, 1: 113. 1854 — Fig. 8

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Fig. 8

Marasmius congregatus (JO61). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. cheilocystidia: e1. Siccus-type broom cells, e2. smooth cells; f. pileipellis: f1. Siccus-type broom cells, f2. smooth cells; g–h. stipitipellis: g. caulocystidia, h. Amyloflagellula-type broom cells. — Scale bars: a = 8.6 mm, b–h = 10 μm.

≡ Chamaeceras congregatus (Mont.) Kuntze, Revis. Gen. Pl. (Leipzig) 3: 455. 1898.

Pileus 5–66 mm diam, hemispheric to almost globose when young, hemispheric to convex when mature, smooth, translucent-striate, slightly sulcate at the margin, centre flat to somewhat depressed, sometimes umbonate, margin incurved to plane, sometimes revolute, edge regular to crenate, frequently lacerate; light orange (N₀₀Y₈₀M₃₀) to brownish orange (N₂₀Y₉₀M₅₀) when immature, becoming paler, whitish cream (N₀₀Y₁₀M₀₀ to N₀₀Y₀₀M_10–20) or pale yellow (N₀₀Y_30–50M_00–10 to N₁₀Y₇₀M₂₀), with light brownish yellow sulci and centre (N₀₀Y₅₀M₁₀ to N₂₀Y₆₀M₄₀), or completely light orange (N₀₀Y₅₀M_10–20); membranous, fragile, context thin (< 1 mm); surface glabrous, humid to moist, semitranslucent or pellucid, especially near the margin, dull at the central disc, papyraceous, hygrophanous. Lamellae free to subfree when mature, adnate to adnexed when very young, subclose to seemingly close because of several lamellulae, L = 16–20, unequal, regular, rarely intervenose, or forked, l = 4, very narrow, smooth, translucent to opaque, white to whitish cream (N₀₀Y₁₀M₀₀), edge even, concolorous with the lamellar faces as well as the hymenium between them. Stipe 15–85 × 1–4 mm, central, cylindrical, generally expanded above and below, or entirely equal, regular; cartilaginous with a harder, chitinous base, hollow; apex and middle concolorous with the lamellae, fading to yellow or orangish (N₁₀Y₉₉M_10–20) or brownish orange (N₂₀Y₇₀M_40–50), reaching orangish brown (N₅₀Y₉₉M₇₀) or dark reddish brown (N_70–80Y_90–99M_70–80) at the base; surface glabrous, but almost imperceptibly pubescent at the base, smooth, with a bright silky sheen; basal mycelium abundant, white to yellow, strigose to almost tomentose.

Basidiospores 5–9 × 2.5–4 μm (x_rm = 7–7.7 × 2.5–4 μm; x_mm = 7 (± 0.4) × 3.4 (± 0.2) μm; Q_rm = 2.1–2.5, Q_mm = 2.1 (± 0.3); n/s = 30, s = 3), somewhat oblong, clavate, subfusoid, or lacrimoid, smooth, hyaline, thin-walled, inamyloid. Basidia 17.3–26.8 × 4–5.6 μm, clavate or cylindrical, slender, hyaline, thin-walled, inamyloid, 2–4 sterigmata. Basidioles 14–24 × 3.5–5.4 μm, cylindrical, clavate or apically tapered, hyaline, inamyloid, thin-walled. Pleurocystidia absent. Cheilocystidia similar to the elements of the pileipellis, consisting of two cellular types: 1) Siccus-type broom cells, main body 13.8–23.8 × 4.4–8.8 μm, clavate to turbinate, cylindrical or irregular in outline, hyaline, with thin to moderately thick walls; setulae erect, apical, short to elongate, 3.8–8 × 0.9–1.3 μm, cylindrical, somewhat dendriform, regular to irregular in outline, simple or branched, solid, with an acute to obtuse apex; 2) Globulares-type smooth cells, 12.5–20 × 7–11.3 μm, pyriform to vesiculose, hyaline, thin-walled, inamyloid. Lamellar trama dextrinoid, irregular, formed by interwoven hyphae, cylindrical, 2.3–10 μm diam, regular, branched, hyaline, thin-walled, smooth. Pileus trama strongly dextrinoid, irregular, interwoven hyphae, cylindrical, 1.8–10.8 μm diam, some segments inflated, branched, very hyaline, thin-walled. Pileipellis hymeniform, composed of two cellular types: 1) Siccus-type broom cells, main body 14.0–25.7 × 6–14.4 μm, clavate to vesiculose, turbinate to pyriform, thin-walled at the base and thick-walled at the apex, hyaline; setulae erect, apical, very short to moderately elongate, 2.2–9 × 0.8–1.5 μm, rare to abundant, cylindrical or conical, regular to irregular in outline, rarely branched, solid, with an acute to obtuse apex, some vesiculose; 2) Globulares-type smooth cells, 14.7–26.7 × 7.6–13 μm, pyriform to vesiculose or balloon-shaped, hyaline, thin-walled. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, somewhat broad, 5–15.2 μm diam, regular in outline, branched, very pale yellowish in KOH, but the hyphae of the stipe base are yellowish brown, with moderately thick walls, septate; internal hyphae very similar to those of the cortex, but narrower, more hyaline, and thin-walled. Stipitipellis with rare caulocystidia, inamyloid, small, 5–21.3 × 2.8–6.3 μm, cylindrical, clavate or with a tapered and acute apex, or even inflated like a vesicle, hyaline, with walls moderately thick, rising from the cortical hyphae. Amyloflagelulla-type broom cells also present, even more rare than the caulocystidia, with main body 6.3–20 × 2.5–5 μm, cylindrical to clavate, hyaline, thin-walled; setulae filiform, thin, elongate, flagellum-like, 7.5–11.3 × 0.5–0.6 μm, solid, regular, flexible, curved, some branched. Clamp connections present in all tissues.

Habit & Substrate — Gymnopoid, scattered, or frequently gregarious or caespitose, on dried leaves and barks of twigs, or rotten wood of dicotyledonous plants.

Distribution — The species was originally described from French Guiana and it was previously recorded from São Paulo, Brazil (Puccinelli & Capelari 2009).

Specimens examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 18 Mar. 2010, J.J.S. Oliveira JO61 (SP 445415!); ibid., J.J.S. Oliveira JO62 (SP 445416!); ibid., 21 Apr. 2010, J.J.S. Oliveira & J.F. Santos JO100 (SP 445426!); ibid., 25 Apr. 2010, J.J.S. Oliveira & J.F. Santos JO122 (SP 445433!); ibid., 9 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO307 (SP 445482!); São Paulo City, Parque Estadual das Fontes do Ipiranga, 1 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO328 (SP 445495!); ibid., J.J.S. Oliveira & F. Karstedt JO332 (SP 445498!); ibid., 3 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO349 (SP 445779!); ibid., J.J.S. Oliveira & F. Karstedt JO354 (SP 445771!); ibid., 2 Feb. 2012, J.J.S. Oliveira JO468 (SP 445553!); ibid., 16 Dec. 2014, J.J.S. Oliveira & G.F. Silva JO556 (SP 466008!); Parque Estadual da Cantareira, Núcleo Engordador, 30 Jan. 2012, J.J.S. Oliveira & V. Motato-Vásquez JO463 (SP 445498!); ibid., J.J.S. Oliveira & M. Capelari JO478 (SP 446040!); ibid., J.J.S. Oliveira & M. Capelari JO511 (SP 446039!); ibid., 16 Feb. 2012, J.J.S. Oliveira & M. Capelari JO512 (SP 446038!); ibid., J.J.S. Oliveira & M. Capelari JO513 (SP 446037!); Iporanga City, Parque Estadual Turístico do Alto Ribeira, Lageado, 2 Mar. 2012, J.J.S. Oliveira & D.E. Desjardin JO531 (SP 446036!).

Notes — Marasmius congregatus was described by Montagne (1854) from French Guiana. It was later recorded in Sri Lanka by Petch (1948) who considered Cantharellus elegans as a synonym of the species. However, in a review of Marasmius pellucidus, Wannathes et al. (2004) noted that what was previously determined as M. congregatus by Petch (1948) actually corresponded to M. pellucidus and, consequently, synonymized C. elegans under M. pellucidus. In the same study, Wannathes et al. (2004) analyzed the holotype of M. congregatus and provided a modern but limited description (for instance, no basidiospores were found in the holotype), recognizing the taxon as an independent species in Marasmius sect. Sicci subsect. Siccini ser. Leonini. Wannathes et al. (2004) also commented that, perhaps, M. congregatus may have been ignored by Dennis (1951b, c) and Singer (1965, 1976) in their studies in South America, and might have considered M. congregatus as a nomen dubio. Based on a type study by Wannathes et al. (2004), however, M. congregatus is a singular species based on the type material labelled ‘Leprieur # 1081 (PC)’ from French Guiana.

Puccinelli & Capelari (2009) identified some specimens of M. congregatus from São Paulo based on the description provided by Wannathes et al. (2004). However, the specimens from São Paulo have short caulocystidia (16–36 mm long) on the stipe base, which were not observed in the holotype (Wannathes et al. 2004). The collections herein examined, also from São Paulo, agree with those studied by Puccinelli & Capelari (2009), especially by having rare and short caulocystidia on the stipe base, but with shorter basidiospores (up to 9 vs 10 μm). The presence or absence of caulocystidia is not conclusive in the holotype, probably because its stipe base is damaged (D.E. Desjardin, pers. comm. apud Puccinelli & Capelari 2009), and, perhaps, because of the short size of these cystidia. Also concerning the stipitipellis, the Amyloflagellula-type broom cells found are similar to those found in M. sullivantii (f. 30 in Gilliam (1976)) and we have interpreted it as an occasional characteristic.

Marasmius pseudoniveus (Singer 1976) is a closely related species, mostly differing by having a distinctly sulcate, pure white pileus; remarkably pubescent-pruinose stipe; larger basidiospores up to 10.3 μm long; and abundant, conspicuous and larger caulocystidia, ranging from 20–60(–100) μm long. The phylogenetic placement of M. congregatus in this study is consistent in the Datasets 2.1 and 3 trees (Fig. 1b, ​,2),2), appearing as sister to M. pseudoniveoaffinis, within the subclade /corrugatus, with strong support. This relationship is further supported by the similar morphology of these two species.

Marasmius pseudoniveoaffinis J.S. Oliveira, sp. nov. — MycoBank MB811131; Fig. 9

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Fig. 9

Marasmius pseudoniveoaffinis (M. Capelari & L.A.S. Ramos 4536). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. cheilocystidia; f. pileipellis: Siccus-type broom cells and smooth cells; g. caulocystidia. — Scale bars: a = 10 mm, b–g = 10 μm.

Holotype. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, on twigs and dried branches of dicotyledonous tree, 2 July 2009, M. Capelari & L.A.S. Ramos 4536 (SP 445583!).

Etymology. Refers to the affinity with Marasmius pseudoniveus in morphology.

Pileus up to 65 mm diam, convex to plane, sulcate-striate or irregularly sulcate, often pure white. Lamellae adnate to free, distant (9–16), with lamellulae inserted (3), regular or interveined-anastomosed. Stipe 8–70 × 1–3 mm, cylindrical, brown, with a strigose basal mycelium. Basidiospores 7–10.4 × 3–4 μm. Cheilocystidia hyphoid, elongate, irregularly cylindrical, clavate, often forked or ramose. Pileipellis hymeniform, made up by both Siccus-type and Globulares-type cells. Caulocystidia cylindrical to setoid. On decaying twigs or branches of dicotyledonous trees.

Pileus 4.5–65 mm diam, convex tending to plane, sulcate-striate or slightly and irregularly sulcate, centre flat or shallowly depressed, margin incurved tending to straight, edge regular, sometimes lacerate; most often completely pure white, but sometimes cream (N₀₀Y₁₀M₀₀), centre sometimes yellowish brown (N₂₀Y_50–60M_20–40); membranous, context thin (< 1 mm); surface glabrous, dry, a little translucent to opaque, papyraceous to subvelutinous, non-hygrophanous. Lamellae adnate to sinuate, sometimes free, distant, L = 9–16, equal or unequal, l = 3, regular or interveined-anastomosed, forked or almost reticulate, faces and edges concolorous with the pileus. Stipe 8–70 × 1–3 mm, central, cylindrical, sometimes a little broader upward to the apex, tough, cartilaginous to horny or chitinous downwards from the middle to the base, hollow; apex white to cream, grading from yellowish or orange brown (N₂₀Y₇₀M₅₀ to Y₉₀M₅₀C₂₀) to dark chestnut brown (N₈₀Y₉₉M₇₀ or Y₉₀M₉₉C₈₀) at the base; surface pruinose to pubescent at the base, opaque; with a well-developed, strigose, cream to whitish brown basal mycelium.

Basidiospores (6.3–)7–10.4(–11.3) × (2.5–)3–4 μm (x_rm = 8.3–9.1 × 3.2–3.8 μm; x_mm = 8.7 (± 0.3) × 3.5 (± 0.3) μm; Q_rm = 2.3–2.7, Q_mm = 2.5 (± 0.2); n/s = 30, s = 4), subellipsoid, lacrimoid to subclavate, hyaline, thin-walled, inamyloid. Basidia 18–22.5(–25) × 6.3–7.5 μm, clavate, hyaline, 2–4 sterigmata. Basidioles 18.8–22.5 × 5–7(–8.8) μm, clavate, hyaline, thin-walled. Pleurocystidia absent. Cheilocystidia inconspicuous, hyphoid, elongate, irregularly cylindrical to clavate, without diverticula, sometimes mucronate, but more often forked, rarely ramose; main body 12.5–52.5 × 3.8–9 μm, hyaline, inamyloid, thin-walled; some with appendices or apical, broad branches with lumen, 3.8–17.3 × 1.8–3.3 μm. Lamellar trama strongly dextrinoid, irregular, hyphae interwoven, cylindrical, 1.3–8.8 μm diam, smooth and regular in outline, some branched, hyaline, thin-walled. Pileus trama strongly dextrinoid, irregular, hyphae interwoven, cylindrical, 2–17.5 μm diam, abundantly septate, similar to those in the lamellar trama in the other aspects. Pileipellis hymeniform, composed of two cellular types: 1) Siccus-type broom cells, main body 10–30.6 × 7.5–16.3(–18) μm, pyriform to cylindrical-clavate, hyaline, thin-walled; setulae apical, erect, elongate to very short, 2–18 × 1–2.5 μm, cylindrical, digitiform or verruciform, regular or irregular in outline, abundant or scarce, solid or with lumen, apex acute to obtuse; 2) Globulares-type smooth cells, more numerous than the first type at the mature basidiomata, 16.3–26.3 × 12.5–17.5 μm, pyriform or balloon-shaped, vesiculose to clavate, or irregular in outline, hyaline, apical portion with thicker walls, dextrinoid. Stipe trama strongly dextrinoid, cortical hyphae parallel, cylindrical, 2.5–17.5 μm diam, regular in outline, yellowish, thick-walled; internal hyphae and those of the stipe apex completely hyaline, thin- to thick-walled, smooth. Stipitipellis containing caulocystidia cylindrical to setoid, some irregular in outline, short to moderately elongate, 16.3–62.5 × 3.8–8.8 μm, some branched, with a rounded apex, hyaline, more numerous and longer than those of the stipe apex, dextrinoid solely in the base, otherwise inamyloid, thick-walled. Clamp connections present in almost all tissues, inconspicuous to conspicuous, not observed in the cortical stipe trama.

Habit & Substrate — Gymnopoid, scattered, on twigs and dried branches of dicotyledonous tree.

DNA barcode — GenBank KP635189 (ITS1-5.8S-ITS2) and KP635144 (LSU).

Distribution — This new species is known from Brazil and Costa Rica (Ovrebo 1996).

Additional specimens (paratypes) examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 18 Mar. 2010, J.J.S. Oliveira JO60 (SP 445414!); ibid., 21 Mar. 2010, J.J.S. Oliveira JO70 (SP 445421!); ibid., 8 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO321 (SP 445917!). – Costa Rica, Heredia Province, La Selva Biological Reserve, Camino Experimental Sul, as Marasmius pseudoniveus var. pseudoniveus Singer, 17 Mar. 1991, Ovrebo 3090 (CSU!).

Notes — Marasmius pseudoniveoaffinis is similar to Maras-mius pseudoniveus (Singer 1965, 1976), especially in the colour of the pileus, macro features of the stipe, size of basidiospores, composition of the pileipellis and the elements of the stipitipellis. However, M. pseudoniveus differs by having a hygrophanous pileus (22–45 mm diam), subclose to almost crowded, regular lamellae instead of distant, often interveined-anastomosed lamellae, and by the cheilocystidia in form of the typical Siccus-type broom cells rather than irregular, elongate, clavate or filamentous, to often forked or ramose cheilocystidia.

Marasmius pseudoniveus var. amylocystis (Singer 1976) is also similar to M. pseudoniveoaffinis mainly by the similar elements at the lamellar edge. However, M. pseudoniveus var. amylocystis differs especially by having shorter basidiospores (7.7–8.7 μm long), by having dextrinoid cheilocystidia, also present at the lamellar faces like pleurocystidia; and by having dermatocystidia in the pileipellis similar to those of the hymenium. Marasmius pseudoniveoaffinis is also similar to M. cohortalis in the pileus pigmentation and by having intervenose-anastamosed lamellae, but differs in having somewhat longer basidiospores (7–10.4 vs 7–8.3 μm, respectively), the presence of peculiar hyphoid cheilocystidia and the pileipellis composed of both Siccus-type broom cells and Globulares-type smooth cells.

The collection from Costa Rica (Ovrebo 3090) identified as M. pseudoniveus var. pseudoniveus by Ovrebo (1996) was also examined and this material has the morphological characteristics completely in concordance with M. pseudoniveoaffinis, especially regarding the cheilocystidia, and is re-determined as conspecific herein. This information shows that this species presents a wide distribution in the Neotropics, at least throughout the zone covered by Atlantic and Amazon forest and their connections between them and with tropical forest in Central America.

3 — Marasmius sect. Globulares Kühner emend. Antonín & Noordel. group Sicci Singer ser. Haematocephali Singer

[Sect. Sicci sensu Singer (1986)]

Marasmius altoribeirensis J.S. Oliveira, sp. nov. — MycoBank MB811132; Fig. 10, ​,1111

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Fig. 10

Marasmius altoribeirensis (JO532). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. pleurocystidia; f. cheilocystidia. — Scale bars: a = 12.8 mm, b–f = 10 μm.

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Fig. 11

Marasmius altoribeirensis (JO532). a. Siccus-type broom cells of the pileipellis; b. broom cells of the stipitipellis. — Scale bars: 10 μm.

Holotype. Brazil, São Paulo States, Iporanga City, Parque Estadual Turístico do Alto Ribeira, on rotten dicotyledonous debris, 2 Mar. 2012, J.J.S. Oliveira JO532 (SP 445579!).

Etymology. The epithet refers to Alto Ribeira (São Paulo, Brazil), the name of the area where the holotype was collected.

Pileus 23 mm diam, convex, smooth, slightly sulcate-striate, olive brown. Lamellae free, subdistant (11), with lamellulae (3), pale cream. Stipe 58 × 2 mm, cylindrical, ochraceous brown, with a tomentose basal mycelium. Basidiospores 6.3–10 × 3.4–5.6 μm, ellipsoid to tear-shaped. Pleurocystidia 30–55 × 4.4–8.8 μm, not projecting, vermiform, ventricose, monilioid or capitate. Pileipellis hymeniform, with brownish Siccus-type broom cells bearing apical, elongate setulae. Stipitipellis containing Siccus-type broom cells.

Pileus 23 mm diam, convex, smooth, or very slightly sulcatestriate, centre flat or somewhat umbonate, margin incurved, edge regular; centre dark-brown (N₉₀Y₉₀M₆₀), becoming olivaceous brown (N₆₀Y₉₉M₅₀); membranous, context cream, thin (≤ 1 mm); surface glabrous, somewhat micaceous, dry, dull, subvelutinous, non-hygrophanous. Lamellae free, subdistant, L = 11, unequal, regular, l = 3, opaque, cream (N₀₀Y₁₀M₀₀); edge even, concolorous as well as the hymenium between the lamellae. Stipe 58 × 2 mm, central, cylindrical, equal, semicartilaginous to cartilaginous, hollow; apex concolorous with the lamellae, becoming ochraceous brown (N₆₀Y₉₉M₅₀); surface glabrous or slightly pruinose at the apex, smooth, with a silky shine; basal mycelium white, tomentose.

Basidiospores 6.3–10 × 3.4–5.6 μm (x_m = 8 (± 2) × 4.5 (± 0.6) μm, Q_m = 1.8 (± 0.2), n = 32), obovoid to suboblong, ellipsoid or lacrimoid, smooth, hyaline, thin-walled, inamyloid. Basidia 30.6–35 × 6.3–7.5 μm, clavate, noticeably elongate, hyaline, thin-walled, inamyloid, few observed. Basidioles 27.5–38.8 × 5–8(–10.3) μm, clavate, noticeably elongate, smooth, hyaline, inamyloid, thin-walled. Pleurocystidia 30–55 × 4.4–8.8 μm, more frequent near the lamellar edge, inconspicuous, deeply rooted in the subhymenium, not projecting much beyond the basidia/basidioles, generally irregular in outline, worm-shaped, sometimes ventricose, monilioid near the apex, or capitate, smooth, somewhat opaque, inamyloid. Cheilocystidia similar to the Siccus-type broom cells in the pileipellis, or completely hyaline, rare, main body 10–21.3 × 3.8–8.8 μm, cylindrical to clavate, turbinate, thin- or thick-walled; setulae apical, erect, generally elongate, 6.3–17.5 × 0.8–1.3 μm, cylindrical, filiform, setoid, regular in outline, simple, pale, solid. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical to inflated, 5–16.3 μm diam, regular or irregular in outline, branched, hyaline, smooth, thin-walled. Pileus trama dextrinoid, irregular, hyphae interwoven, cylindrical to inflated, 2.5–17 μm diam, similar to those of the lamellar trama. Pileipellis hymeniform, composed of Siccus-type broom cells, appearing brown in mass, some hyaline when isolated, main body 12.5–25 × 3.8–13.8 μm, cylindrical, clavate, frequently turbinate, sometimes branched, walls slightly thick at the apex, but becoming thinner toward the base; setulae apical, erect, generally very elongate, 6.3–18.3 × 0.6–2 μm, cylindrical, thin, filiform, setoid, simple, brown, apex acute or slightly obtuse. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, 4.4–12.5 μm diam, regular in outline, melleous or browner, smooth, walls slightly thick; internal hyphae hyaline, thin-walled, 2.5–13.8 μm diam. Stipitipellis with Siccus-type broom cells, more often present at the apical portion of the stipe, main body generally reduced, 5–13.8(–18.8) × 2.5–12.5 μm, many times irregular in outline, thick-walled, hyaline to pale brown; setulae very elongate, 3.8–23.8 × 0.9–2.5 μm, cylindrical, filiform to thick setoid, appearing tough, solid or with lumen, regular in outline, simple. Clamp connections present in all tissues.

Habit & Substrate — Gymnopoid, solitary, on rotten dicotyledonous debris.

DNA barcode — GenBank KP635204 (ITS1-5.8S-ITS2) and KP635158 (LSU).

Notes — Marasmius altoribeirensis is one of the few species of Marasmius combining olivaceous pileus and pleurocystidia. It also has broom cells of the pileipellis and stipitipellis with very long apical setulae. Few species with olivaceous pileus are known in Marasmius (Antonín 2007). One example is M. brunneoolivacens, which strongly differs from the new species by the much more elongate basidiospores (12–14 μm) and by the presence of two types of pleurocystidia, one in form of Siccus-type broom cells and the other in form of cylindrical to clavate cells, sometimes with lobes or constrictions, but not diverticulate (Wannathes et al. 2009). Marasmius elaeocephalus sensu Antonín (2007) is another species with olive-brown pileus very similar to M. altoribeirensis, but differs by having thinner basidiomata, smaller pileus (6–14 mm diam), slender stipe, larger basidiospores (10.8–13.8 μm long), refractive, slightly shorter pleurocystidia (19–47 μm long), and not so elongate setulae at the apex of the broom cells.

In the trees recovered, M. altoribeirensis showed an unsupported relationship with M. spegazzinii (Fig. 2), within the strongly supported /confertus clade. These two species share characteristics such as a firm (not thin-membranous) pileus and the presence of medium-sized pleurocystidia.

Marasmius anomalus Lasch ex Rabenh., Klotzsch. Herb. Vivum. N° 1806, pl. 17, f. 97. 1854. (non Marasmius anomalus Peck 1872) — Fig. 12

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Fig. 12

Marasmius anomalus (JO346). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. pleurocystidia; f. cheilocystidia; g. Siccus-type broom cells of the pileipellis. — Scale bars: a = 10.8 mm, b–g = 10 μm.

Pileus 1.4–11.7 mm diam, conical to campanulate, or hemispheric, later convex to plane, centre smooth, flat or somewhat umbonate, smooth in the youngest basidiomata, becoming radially striate or slightly sulcate and sometimes rugulose with age; margin incurved to decurved, edge entire or crenate; centre brownish orange (N₄₀Y₉₀M₆₀), dark orange or almost ochraceous (N₄₀Y₉₉M₆₀), remaining the same or becoming orange (N₁₀Y_80–90M₄₀), or paler orange (N₁₀Y₆₀M₃₀) toward the margin with age; membranous, context thin (< 1 mm); surface glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae free, subfree to adnexed, subdistant to distant, L = 12–14, equal, regular, l = 0, opaque, smooth, white to cream (N₀₀Y₁₀M₀₀), edge even, cream to finely concolorous with the pileus as well as with the hymenium between the lamellae. Stipe 9–38 × 0.2–0.7 mm, central, cylindrical, thin or filiform, equal, simple; tough, cartilaginous to chitinous, hollow; apex concolorous with the lamellar faces, grading to yellowish brown (N₅₀Y₉₉M₄₀), then dark reddish brown (N_80–90Y₉₉M_50–60) or black at the base; surface glabrous, smooth, with a silky shine; strigose basal mycelium white to cream, scarce (visible only under good lens), or stipe subinsititious.

Basidiospores (11.6–)12.5–17.5 × 3–4.5(–5) μm (x_rm = 15.3–15.7 × 3.6–3.8 μm; x_mm = 15.3 (± 0.4) × 3.7 (± 0.1) μm; Q_rm = 4–4.2; Q_m = 4.2 (± 0.1); n/s = 30, s = 3), oblong, clavate, smooth, hyaline, thin-walled, inamyloid. Basidia 18.8–26.3 × 8.4–12 μm, clavate, smooth, hyaline, 3–4 sterigmata, thin-walled, inamyloid. Basidioles 18.8–27.5 × 5.6–10 μm, broadly clavate in general, some tapering upward to the apex, tending to have a acute apex, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia 31.3–46.3 × 7.5–9.4 μm, inconspicuous in the hymenial layer, slightly projecting above the basidia, conspicuous when isolated, poorly developed, clavate, frequently capitate, with a small apical vesicle, or mucronate, with slight to strong constrictions upward to the apex, sometimes with a tapered apex, smooth, somewhat opaque, thin-walled, inamyloid. Cheilocystidia very similar to the Siccus-type broom cells in the pileipellis, with the same colour, abundant, main body 8.8–17.5 × 5.6–7.5 μm, clavate to slightly turbinate, sometimes ventricose, thin-walled, hyaline; setulae apical, erect, 1.3–6.3 × 0.6–1 μm, cylindrical, filiform, thin, digitiform, regular in outline, pale orange, simple, solid, apex slightly obtuse or frequently acute. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2–9.4(–20) μm diam, regular or irregular in outline, with some inflated segments, branched, smooth, hyaline, thin-walled. Pileus trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2.3–12.5 μm diam, similar to those of the lamellar trama. Pileipellis hymeniform, composed of abundant Siccus-type broom cells, orangish to melleous when in group, main body 12.5–20 × 4.4–13.8 μm, clavate, turbinate, ventricose, thin-walled, hyaline; setulae apical, erect, 1.5–5(–7.5) × 0.5–1 μm, cylindrical, thin, finger-like, simple, regular in outline, yellowish, solid, apex slightly obtuse or almost acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, 2–8.8 μm diam, regular in outline, melleous, yellowish to pale brown, smooth, thick-walled; internal hyphae hyaline, thin-walled, 2.5–8.8 μm diam. Clamp connections present in almost all tissues, but absent in the cortical stipe trama.

Habit & Substrate — Marasmioid, gregarious or scattered, on rotten twigs of dicotyledonous tree.

Distribution — The species was described from Driesen, Germany, and recorded from Austria, Czech Republic, French, Italy, Netherlands, Romania, Slovakia and Spain in Europe (Antonín & Noordeloos 2010); Algeria, Morocco (Antonín & Noordeloos 2010) and Cameroon in Africa (Douanla-Meli & Langer 2008); and from Argentina and Chile in South America (Singer 1976). In Brazil, M. anomalus has previously been recorded from Paraná State (De Meijer 2001) and this is the first record from São Paulo State.

Specimens examined. Brazil, São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, 1 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO340 (SP 445777!); ibid., 3 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO346 (SP 445506!); ibid., 23 Mar. 2011, J.J.S. Oliveira & P.O. Ventura JO361 (SP 445774!).

Notes — Singer (1976) studied the holotype (Lasch - FH) from Driesen and confirmed it to be conspecific with collections from Tucumán and Cordoba in Argentina and Santiago in Chile. Antonín & Noordeloos (2010) examined collections from Austria, Czech Republic, French, Germany, Italy, the Netherlands, Romania, Slovakia and Spain, and provided a description of the species with much larger basidiospores ((12–)16–22(–24) × (3–)4–5(–6) μm), very elongate basidia and basidioles (up to 42 μm long), shorter pleurocystidia (36–60 μm long) and some inflated hyphal segments in the lamellar and pileus trama. Their description differed from that published by Singer (1976) and the collections examined by us and may be due to a mixture of collections of different species. Antonín & Noordeloos (2010) also listed other inconsistencies in the literature such as larger pileus reaching 20 mm diam (Battetta 1934, Clémençon 1982), more numerous lamellae ranging from 10–20 (Noordeloos 1987, Singer 1965, 1976), smaller basidia measuring from 28–30 μm (Battetta 1934, Singer 1976).

The material herein examined are concordant in most morphological aspects with those collections studied by Singer (1976), in others with collections studied by Antonín & Noordeloos (2010), or with both. The dimensions of the basidiospores and almost all other characteristics are consistent with those of Singer (1976) and with the protologue, especially regarding basidiospore dimensions, and differs only by the smaller number of lamellae (12–14 vs 10–20) and by shorter pleurocystidia (31.3–46.3 vs 30–83 μm). On the other hand, the number of lamellae and the size of pleurocystidia are in agreement with Antonín & Noordeloos (2010). However, according to these latter authors, the basidia/basidioles as well as the basidiospores are more elongate. Nonetheless, the specimens from São Paulo fit into the species in a ‘broad concept’, considering a large range in the basidiospores size. Furthermore, colour pictures of a specimen provided by Antonín & Noordeloos (2010) show that the material examined by them resembles ours. The collection from Paraná has no morphological description in published papers, and only has been cited (in tables) in a survey of macrofungi in the state of Paraná (De Meijer 2001). Unfortunately, the material could not be examined in this study.

Desjardin (1989) established the lectotype of M. anomalus on the isotype ‘Germany, Driesen, Lasch n° 1806 (FH)’, providing a full morphological description which is mostly concordant with this present description, supporting this identification. The only distinction is that the lectotype presents non-marginate lamellae. Nonetheless, it is still not completely certain if the specimens from South America (here we include those specimens examined in Singer (1976)) are actually M. anomalus described from Germany and widely distributed across Europe and, thus, a molecular comparison should be done. Currently, the species seems to be globally distributed in a most general morphological sense, with the obvious exception of arctic, subarctic, arid and subarid areas. Singer (1976) argued that this is not a typical Neotropical species, but has entered the American subtropics at least in the Southern Hemisphere.

Unfortunately, only ITS sequence could be obtained from JO346 M. anomalus while DMC 011 M. anomalus, a specimen collected from Cameroon (Africa), has only nLSU sequence (EF160086) deposited in GenBank, and thus the two strains could not be compared. No sequence from European collections (especially from Germany) representing the species is available for comparison so far. However, JO346 herein representing the singerian concept of a ‘neotropical M. anomalus’ is shown in Fig. 1c as potentially conspecific with NW285 M. hypophaeus (Wannathes et al. 2009). Both JO346 and NW285 specimens are morphologically conspecific, with exception of the broom cells also present on the lamellar faces along with non-setulose, medium-sized pleurocystidia (41–48 μm) found in NW285. Moreover, the collection NW285 seems to correspond to the description of M. anomalus in Singer (1976) and Desjardin (1989), despite sharing some similarities with M. hypophaeus.

Marasmius luteoolivaceus J.S. Oliveira, sp. nov. — MycoBank MB811133; Fig. 13

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Fig. 13

Marasmius luteoolivaceus (JO524). a. Basidiomata; b. basidiospores; c. basidioles; d. pleurocystidia; e. cheilocystidia; f. Siccus-type broom cells of the pileipellis. — Scale bars: a = 13.4 mm, b–f = 10 μm.

Holotype. Brazil, São Paulo State, Iporanga City, Parque Estadual Turístico do Alto Ribeira, Núcleo Santana, on dried leaves of dicotyledonous tree, 29 Feb. 2012, J.J.S. Oliveira & D.E. Desjardin JO524 (SP 445574!).

Etymology. From the Latin luteo = yellow; olivaceus = olivaceous, referring to the pileus colour.

Pileus 1.5–12 mm diam, hemispherical to convex, shallowly sulcate, umbonate, yellowish olive to greyish yellow. Lamellae free, subdistant (16–18). Stipe 8.5–38.5 × 0.5–1 mm, reddish brown, with a strigose basal mycelium. Basidiospores 9.3–12.5 × 3–3.8 μm, clavate to subfusoid. Pleurocystidia 23.8–40 × 3.8–6.5 μm, clavate, slightly ampullaceous, hyaline. Lamellae and pileus trama dextrinoid, irregular. Pileipellis hymeniform, composed of brownish Siccus-type broom cells.

Pileus 1.5–12 mm diam, hemispheric to convex, slightly sulcate, centre shallowly to strongly umbonate, margin decurved, edge regular to wavy; pure grey to olivaceous grey (N₈₀Y_40–50M_10–20) when young, then pale olivaceous brown (N₇₀Y_90–99M_00–20), yellowish olive (N₅₀Y_90–99M_10–20) or greyish yellow (N₄₀Y_40–60M₁₀) when mature, fading to pale brownish yellow (N₅₀Y₈₀M₄₀) toward the margin, with a dark olive centre (N₈₀Y₉₀M₀₀) or dark olivaceous brown (N₇₀Y₉₉M₁₀); membranous, context cream, thin (< 1 mm); surface glabrous, dry, dull, papyraceous to subvelutinous, non-hygrophanous. Lamellae free, subclose to subdistant, L = 16–18, equal, regular, l = 0, opaque, smooth, pale cream (N₀₀Y₁₀M₀₀), edge even, concolorous with the lamellar face or finely concolorous with the pileus, as well as with the hymenium between lamellae. Stipe 8.5–38.5 × 0.5–1 mm, central, cylindrical or filiform, equal, semicartilaginous to chitinous, hollow; apex concolorous with the lamellar face, becoming yellowish brown (N₄₀Y₇₀M₄₀) or orange (N₆₀Y₉₀M₃₀) and reddish brown (N₈₀Y₉₉M₇₀) toward the base; surface glabrous, smooth, shiny; with a yellowish, strigose basal mycelium.

Basidiospores 9.3–12.5 × 3–3.8 μm (x_m = 11.2 (± 0.9) × 3.3 (± 0.3) μm, Q_m = 3.4 (± 0.4), n = 30), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 17.5–25 × 4–7 μm, clavate to tapered at the apex, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia 23.8–40 × 3.8–6.5 μm, generally cylindrical-clavate, clavate, sometimes tending to ampullaceous, or with constrictions at the apical regions, with a somewhat more inflated apex or a more ventricose base, smooth, hyaline, thin-walled, inamyloid. Cheilocystidia forming a sterile, pale yellow lamellar edge, in form of Siccus-type broom cells, hyaline when isolated, main body 11.3–18.8 × 4.8–6.5 μm, cylindrical-clavate to clavate, sometimes turbinate to ventricose, thin-walled; setulae apical, erect, short to somewhat elongate, 0.8–4.6 × 0.6–1.3 μm, cylindrical, digitiform to verruciform, rarely globose or broadened, regular to irregular in outline, hyaline, solid, simple, apex acute to obtuse. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 1.3–10 μm diam, regular in outline, branched, smooth, thin-walled. Pileus trama dextrinoid, irregular, hyphae similar to those of the lamellar trama, 1.3–8.8 μm diam. Pileipellis hymeniform, composed of Siccus-type broom cells, brownish ochre when in group, main body 8.8–20 × 5–8.8(–12.5) μm, clavate, turbinate to ventricose, rarely forked, hyaline, thin- to slightly thick-walled; setulae apical, erect 1.3–7 × 0.5–1 μm, filiform, setiform, cylindrical, thin, simple, regular in outline, generally elongate, sometimes short and verruciform, simple, hyaline to pale yellow, apex acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, regular in outline, 2.5–6 μm diam, smooth, yellowish or chestnut brown to olivaceous in KOH, walls somewhat thick; internal hyphae 1.3–5.6 μm diam, hyaline, smooth, thin-walled. Clamp connections present in almost all tissues, sometimes inconspicuous, not observed in the cortical stipe trama.

Habit & Substrate — Marasmioid, gregarious, on dried leaves of dicotyledonous tree.

DNA Barcode — GenBank KP635182 (ITS1-5.8S-ITS2) and KP635137 (LSU).

Notes — This new species resembles M. elaeocephalus (Singer 1964, Antonín 2007) in the pileus pigmentation and in many other characteristics. However, M. elaeocephalus differs by having many series of lamellulae (2–4), larger basidiospores (10.8–13.8(–14.5) × 3.8–5.8 μm) and the pleurocystidia are often subfusoid or rostrate, and somewhat refractive (Antonín 2007).

Based on the Dataset 3 tree (Fig. 2), M. luteoolivaceus is strongly supported as sister to M. hobbitii within subclade /graminicola.

Marasmius neotropicalis J.S. Oliveira, sp. nov. — MycoBank MB817805; Fig. 14

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Fig. 14

Marasmius neotropicalis (JO69). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. pleurocystidia; f. cheilocystidia; g. Siccus-type broom cells in the pileipellis. — Scale bars: a = 9.2 mm, b–g = 10 μm.

Holotype. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, on dried twigs of dicotyledonous tree, 21 Mar. 2010, J.J.S. Oliveira JO69 (SP 445420!).

Etymology. The epithet refers to ‘Neotropical’.

Pileus 5–20 mm diam, convex to plano-convex, smooth or shallowly sulcate, often bright orange. Lamellae free, subdistant (12–16), with 2–3 series of lamellulae. Stipe 16–34 × 0.8–1.3 mm, brownish orange, with a strigose basal mycelium. Basidiospores 7.5–13.5 × 2.8–4 μm, clavate to subfusoid. Pleurocystidia 27.5–44.5 × 5–7.5 μm, cylindrical, lageniform, clavate, submoniliform, acuminate or capitate. Cheilocystidia in form of broom cells with short nodulose setulae. Trama dextrinoid. Pileipellis hymeniform, composed of Siccus-type broom cells, having setulae (2.5–)4.8–14.3(–17.5) × 0.5–1.6 μm. On dried branches and thin twigs of dicots.

Pileus 5–20 mm diam, convex to plano-convex, smooth or more often sulcate-striate at least on the margin, centre flat or umbonate, margin incurved to decurved, tending to plane, edge regular or slightly undulate; pale brownish orange (N₂₀Y₆₀M₅₀), pale orange, bright orange to frequently darker orange (N₀₀Y_50–80M_10–50, N₁₀Y₅₀M₄₀ to N₁₀Y₆₀M₅₀), sometimes pale yellow (N₀₀Y₃₀M₀₀), centre yellowish brown (N₂₀Y₆₀M₄₀) to orangish brown (N₄₀Y₉₉M₆₀); membranous, context thin (< 1 mm); surface glabrous, dry, sometimes a little translucent, usually dull, subvelutinous, non-hygrophanous. Lamellae free, subdistant, L = 12–16, equal, l = 2–3, rarely forked or slightly reticulate, faces and edges white to pale cream (N₀₀Y₂₀M₀₀). Stipe 16–34 × 0.8–1.3 mm, central, cylindrical, equal, semicartilaginous to slightly chitinous downwards, tough, hollow; apex concolorous with the lamellae or pale yellow (N₀₀Y₅₀M₁₀), becoming yellowish brown (N₃₀Y₈₀M₆₀) or orangish brown (N₂₀Y₆₀M₆₀) in the middle, reaching dark brown (N₄₀Y₉₉M₈₀ to N₉₀Y₉₉M₇₀) near the base; surface glabrous, smooth, opaque or with a somewhat silky shine; basal mycelium strigose, abundant, long-filamentose, cream to yellowish.

Basidiospores 7.5–13.5 × 2.8–4 μm (x_rm = 9.6–11.7 × 3.5–3.6 μm; x_mm = 10 (± 1.1) × 3.6 (± 0.1) μm; Q_rm = 2.7–3.2; Q_mm = 2.7 (± 0.3); n/s = 30, s = 3), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia 18.8–29 × 6–9 μm, clavate to vesiculose, generally with an inflated apex, hyaline, thin-walled, inamyloid, 2–4 sterigmata. Basidioles 18.5–28 × 5.6–9.3 μm, clavate or tapering towards the apex, hyaline, inamyloid. Pleurocystidia 27.5–44.5 × (3.4–)5–7.5 μm, sparse, inconspicuous, not prominently projecting beyond the basidia, some with the basal portion more deeply rooted in the subhymenium, cylindrical, lageniform, clavate to submoniliform, apex acuminate, sometimes capitate or with an apical, vesicle or mucro-shaped capitule, smooth, hyaline, thin-walled, inamyloid, some simply as cystidioles with a fusoid apex. Cheilocystidia numerous, somewhat different from the Siccus-type broom cells of the pileipellis, main body 10.6–20 × 4.5–8 μm, clavate to vesiculose, sometimes lobed, hyaline, thin-walled; setulae apical, erect, short, 1–3.8(–5) × 0.5–1.3 μm, many nodulose or irregularly finger-like to conical, others submonilioid, many branched and vesiculose, coralloid, irregular in outline, hyaline, solid, abundant to scarce, apex acute or mostly obtuse and rounded. Lamellar trama strongly dextrinoid, irregular, hyphae interwoven, cylindrical, 2–11 μm diam, regular in outline, branched, hyaline, thin-walled, smooth. Pileus trama strongly dextrinoid, irregular, hyphae similar to those of the lamellar trama, 2.3–12.5 μm diam. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, pale orange in mass, main body 10–18(–22.5) × 4.6–12.5 μm, clavate to turbinate, somewhat vesiculose, sometimes irregular in outline, forked or branched, hyaline, thin- to slightly thick-walled at the apex; setulae apical, erect, with a moderate length to frequently elongate, (2.5–)4.8–14.3(–17.5 ) × 0.5–1.6 μm, cylindrical, regular in outline, simple or branched, solid, abundant, hyaline to pale yellow, apex acute to slightly obtuse. Stipe trama strongly dextrinoid, cortical hyphae parallel, 3.5–8 μm diam, cylindrical, regular in outline, branched, smooth, yellowish brown, walls moderately thick, septate; internal hyphae similar to those of the cortex, but hyaline, disorganized, 2–9.4 μm diam, thin- to moderately thick-walled. Clamp connections present in all tissues.

Habit & Substrate — Gymnopoid to almost marasmioid, scattered, on dried branches and thin twigs of dicotyledonous tree.

DNA Barcode — GenBank KP635183 (ITS1-5.8S-ITS2) and KP635138 (LSU).

Additional specimens (paratypes) examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 16 Dec. 2009, M. Capelari & L.A. Ramos 4568 (SP 445908!); ibid., 10 Feb. 2010, J.J.S. Oliveira & A.V. Costa JO29 (SP 445909!); ibid., 23 Mar. 2010, J.J.S. Oliveira JO82 (SP 445910!); ibid., 21 Apr. 2010, J.J.S. Oliveira & J.F. Santos JO99 (SP 445912!); ibid., 25 Apr. 2010, J.J.S. Oliveira & J.F. Santos JO118 (SP 445431!); ibid., J.J.S. Oliveira & J.F. Santos JO119 (SP 445913!); ibid., J.J.S. Oliveira & J.F. Santos JO121 (SP 445914!); ibid., 7 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO293 (SP 445473!); ibid., 9 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO325 (SP 445493!).

Notes — Morphologically, M. neotropicalis is very similar to M. suthepensis (Wannathes et al. 2009), differing by having a generally brighter orange pileus; by the nodulose and short setulae at the apex of the completely hyaline Siccus-type broom cells on the lamellar edge, while those in the pileipellis have much more elongate setulae ((2.5–)4.8–14.3(–17.5) vs 3–8 μm); and by the noticeable substrate preference for wood or twigs of dicotyledonous tree instead of leaves.

However, in the Datasets 2.1 and 3 trees recovered (Fig. 1b, ​,2),2), M. neotropicalis appears very close to the M. linderioides complex, within the /graminicola clade. This latter species (type variety) differs by the pileus pigmentation tending to beige, by the very short stipe (1–3 mm vs 16–34 mm long), by the larger basidiospores (10–15 × 3.4–5 μm), by having cheilocystidia with more elongate and regular setulae, and broom cells of the pileipellis having shorter setulae (1.5–4 μm). A pairwise comparison matrix (Table S1) of ITS sequences including M. linderioides, M. neotropicalis and M. suthepensis reveals they are very close, showing a dissimilarity of c. 2–3 % between M. linderioides and M. neotropicalis, and c. 2–3 % between M. neotropicalis and M. suthepensis. The canonical 3 % criterion seems to fail for ITS (Nilsson et al. 2008) to resolve this species complex. This may be due to incomplete data as some sequences are shorter than the expected (due to sequencing artifacts), but also likely shows limitation of the ITS to delimit these species. These three taxa may have recently diverged to form different species and thus there are too few genetic differences found among the taxa of this species group. Interestingly, almost all collections are sympatric where speciation processes are still occurring. The differences in the substrate preferences may also reflect this divergence and serve as a diagnostic character.

Marasmius spegazzinii Sacc. & P. Syd., Syll. Fung. (Abellini) 14: 117. 1899 — Fig. 15

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Fig. 15

Marasmius spegazzinii (JO467). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. pleurocystidia; f. cheilocystidia; g. Siccus-type broom cells of the pileipellis. — Scale bars: a = 20 mm, b–g = 10 μm.

≡ Marasmius balansae Speg., Revista Argentina de Historia Natural 1 (2): 102 (1891).

Pileus 21–45 mm diam, convex to more often becoming plane, tending to revolute, smooth, somewhat sulcate-striate, centre flat or shallowly umbonate, margin straight or somewhat uplifted, edge regular, with some lacerations; centre dark brown (N₉₀Y₉₉M₇₀) or paler (N₈₀Y₉₉M₆₀) or fulvous brown (N₅₀Y₉₉M₈₀), becoming brownish orange or ochre (N₄₀Y₈₀M₅₀) toward the margin; membranous, context cream, thin (< 1 mm); surface glabrous, dry, dull, subvelutinous to velutinous, non-hygrophanous. Lamellae free to subfree, subclose to subdistant, L = 18–26, broad (up to 4 mm), unequal, regular, forked or rarely anastomosed, l = 5 or more, opaque, smooth, whitish cream (N₀₀Y₁₀M₀₀) to more yellowish, edge even, concolorous with the lamellar faces as well as with the hymenium between the lamellae. Stipe 46–85 × 1.5–4 mm, central, cylindrical, sometimes compressed, equal, tough, semicartilaginous to chitinous, hollow; apex concolorous with the lamellar faces, becoming brownish orange (N₅₀Y₉₉M₅₀), dark reddish brown (N₉₀Y₉₉M₇₀) downwards; surface glabrous, smooth, with a silky bright; basal mycelium tinted yellow, strigose.

Basidiospores 7.5–11.3 × 2.5–3.5 μm (x_m = 9.2 (± 1) × 2.9 (± 0.3) μm, Q_m = 3.2 (± 0.3), n = 30), generally oblong, cylindrical, clavate to subfusoid, sometimes subellipsoid, smooth, hyaline, thin-walled, inamyloid. Basidia 22.8–31.3 × 4.6–6.3 μm, cylindrical to clavate, smooth, hyaline, 4 sterigmata, thin-walled, inamyloid. Basidioles (17.5–)20–28.8 × 3.8–7.2 μm, cylindrical to clavate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia 21.3–40 × 6.3–10 μm, abundant, present not only on the lamellar faces, but also at the edge, almost indistinguishable in the hymenial layer, not prominently projecting above the basidia, versiform, frequently clavate to pyriform, some more elongate, ventricose or with 1–3 constrictions near the apex, which is narrower and somewhat monilioid, or only capitate, completely smooth, hyaline, thin-walled. Cheilocystidia in form of Siccus-type broom cells similar to those of the pileipellis, but overall hyaline, rare, main body 11.3–22.5 × 6.3–10 μm, clavate to slightly turbinate, thin-walled; setulae apical, erect, 3.4–13 × 0.6–2 μm, cylindrical to digitiform, regular in outline, simple to branched, hyaline, solid or with lumen, apex frequently acute. Lamellar trama strongly dextrinoid, irregular, hyphae interwoven, cylindrical, 2–10 μm diam, regular in outline, sometimes inflated, branched, smooth, thin-walled. Pileus trama strongly dextrinoid, irregular, or subregular, hyphae resembling those of the lamellar trama, 2.5–16.3 μm diam. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, pale ochre in mass, main body 10–20 × 5–13 μm, more often turbinate, sometimes clavate, lobed or irregular in outline, hyaline, thin- to moderately thick-walled, thinner toward the base; setulae apical, erect, 2.5–11.3 × 0.6–1.5 μm, cylindrical, filiform or setiform, simple to forked, regular in outline, hyaline to pale yellow, apex generally acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, regular in outline, 2.5–12.5 μm diam, smooth or somewhat rough, pale brown to ochre, thick-walled; internal hyphae, 2–15 μm diam, hyaline, smooth, thin-walled. Clamp connections present in almost all tissues, except the cortical stipe trama.

Habit & Substrate — Gymnopoid, scattered, on rotten leaves of dicotyledonous trees.

Distribution — The species was described originally from Guarapí (Paraguay) and subsequently reported from Costa Rica, Peru, Bolivia, Argentina, the United States of America and Tanzania (Singer 1976, Pegler 1977). This is the first record of the species from Brazil.

Specimen examined. Brazil, São Paulo State, São Paulo City, Parque Estadual das Fontes do Ipiranga, 2 Feb. 2012, J.J.S. Oliveira JO467 (SP 445552!).

Notes — The specimen analysed matches consistently the description of M. spegazzinii as described by Singer (1976), differing only by the slightly shorter pleurocystidia (21.3–40 μm vs 27.5–52 μm) and less elongate setulae on the broom cells in the pileipellis. In Pegler (1977), the only relevant difference found was the woody substrate (twigs on the ground) instead of leaves.

The specimen examined is very close to Marasmius spadiceus based on descriptions found in Gilliam (1975, 1976). However, Desjardin (1991) established M. spadiceus as synonym of M. floridanus, the latter previously considered a synonym of M. spegazzinii by Singer (1976). Desjardin (1991) stated that he had not the chance to study the holotype of M. spegazzinii (Balansa 4284, nom. nov. for M. balansae (Spegazzini 1891) non M. balansae (Patouillard 1890)), since it was no longer deposited in the LPS herbarium, and was apparently lost. Therefore, the conspecificity of M. spegazzinii and M. floridanus sensu Singer (1976) was not accepted by Desjardin (1991) until new specimens from Paraguay corresponding to the protologue of M. spegazzinii (Balansa 4284) could be collected.

However, Singer (1976) examined the holotype (Balansa 4284) and a syntype from Guarapi (Paraguay), along with other collections of M. spegazzinii from Costa Rica, Peru, Bolivia and Argentina, and the holotype of M. floridanus deposited in FLAS, Florida, and concluded that the latter species should be a synonym of M. spegazzinii.

The name Marasmius spegazzinii Sacc. & P. Syd. (replacing M. balansae Speg., for it is a competing homonym with the older name M. balansae Pat.) is currently illegitimate (nomen superfluum) in MycoBank, whereas the legitimate name is assigned to Chamaeceras spegazzinii (Kuntze 1898), a nomen novum also for M. balansae Speg. However, Chamaeceras Rebent. ex Kuntze is illegitimate as a synonym of Marasmius Fr. If the priority is for Chamaeceras spegazzinii Kuntze against Marasmius spegazzinii Sacc. & P. Syd., a new combination may be needed based on Chamaeceras spegazzinii Kuntze.

Based on the Dataset 3 tree (Fig. 2), M. spegazzinii appears as an unsupported sister to M. altoribeirensis within the subclade /confertus.

Marasmius suthepensis Wannathes, Desjardin & Lumyong, Fungal Diversity 37: 288. 2009 — Fig. 16

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Fig. 16

Marasmius suthepensis (JO469). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. pleurocystidia; f. cheilocystidia; g. Siccus-type broom cells of the pileipellis. — Scale bars: a = 15.2 mm, b–g = 10 μm.

Pileus 2 mm diam in youngest stages, later 6.8–19 mm diam, convex, hemispheric to campanulate, smooth or striate, centre slightly umbonate with a small umbo, margin incurved to decurved, edge regular; yellowish brown with faint olivaceous hues (N₆₀Y₇₀M₄₀) when immature, becoming yellow (N₁₀Y₉₀M₂₀) to orange yellow (N₁₀Y₉₀M₄₀) or light orange (N₂₀Y₉₀M₅₀) when mature, with centre ochraceous brown (N₄₀Y_70–80M₃₀) to brown (N₅₀Y₉₀M₅₀); membranous, context thin (< 1 mm); surface glabrous, dry, dull, smooth, non-hygrophanous. Lamellae free to adnexed, close, abundant considering the lamellulae, L = 13–23, unequal, regular, l = 3–4, opaque, smooth, whitish cream (N₀₀Y₁₀M₀₀), edge even, concolorous with the lamellar faces, but the hymenium between lamellae is concolorous with the pileus. Stipe 10–68 × 0.6–1 mm, central, cylindrical, equal or sometimes with a broadened apex; tough, semicartilaginous to chitinous, hollow; apex greyish yellow (N₁₀Y₅₀M₂₀) or even concolorous with the lamellae, becoming ochraceous brown or orange (N₃₀Y₇₀M₄₀ to N₄₀Y₆₀M₅₀), reaching dark brown (N₆₀Y₈₀M₇₀) to almost black (N₉₀Y₉₉M₆₀) downward to the base; surface glabrous, smooth, with a silky shine; basal mycelium strigose, scarce to abundant, tinted yellow.

Basidiospores 8.8–14 × 3–5(–5.4) μm (x_rm = 11.3–11.7 × 3.4–4 μm, x_mm = 11.4 (± 0.2) × 3.9 (± 0.1) μm, Q_rm = 2.8–3.1, Q_mm = 3 (± 0.1), n/s = 30, s = 5), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia 20–26.5 × 6–7.5 μm, clavate, hyaline, thin-walled, 2–4 sterigmata, inamyloid. Basidioles 18.8–28.3 × 6.3–7.5 μm, clavate to subacuminate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia 21.3–36.3 × 5–7.5 μm, inconspicuous, poorly developed, smooth, subacuminate, mucronate to capitate, with an evident, vesiculose appendage, or apically capitulate, hyaline, inamyloid. Cheilocystidia abundant, mostly limited to the lamellar edge, but some also dispersed across the face, in form of Siccus-type broom cells, main body 11.3–23.8 × 5–7.8 μm, clavate to turbinate, very similar to those of the pileipellis, thin-walled, hyaline; setulae apical, erect, moderately elongate to rarely short, 2.5–7 × 0.5–1 μm, cylindrical, filiform, hyaline, simple, apex acute. Lamellar trama strongly dextrinoid, irregular, hyphae cylindrical to rarely inflated, 1.5–12.5 μm diam, smooth, branched, hyaline, thin-walled. Pileus trama strongly dextrinoid, irregular, tending to regular near the pileipellis, hyphae cylindrical to inflated, 2–12.5 μm diam, smooth, hyaline, thin-walled. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, pale orange, hyaline when isolated, main body 10–18.8 × 6.3–10 μm, clavate to turbinate, thin-walled, dextrinoid; setulae apical, erect, elongate, 2.3–7.5 × 0.6–1 μm, filiform to cylindrical, very thin, solid, simple, apex acute. Stipe trama strongly dextrinoid, cortical hyphae parallel, cylindrical, 3–10.6 μm diam, regular in outline, smooth, thick-walled, yellowish brown, septate; internal hyphae hyaline, 2.5–11.3 μm diam, thin-walled, branched. Clamp connections present in almost all tissues, but absent in the cortical stipe trama.

Habit & Substrate — Gymnopoid, scattered or gregarious, on petioles of leaves and dried twigs of dicotyledonous tree.

Distribution — Originally described from Thailand (Wannathes et al. 2009) and also found in Príncipe (Grace et al. 2019), it is herein recorded for the first time from Brazil.

Specimens examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 2 Dec. 2009, M. Capelari & L.A.S. Ramos 4556 (SP 446013!); ibid., 14 Jan. 2010, J.J.S. Oliveira & M. Capelari JO3 (SP 446115!); ibid., 23 Mar. 2010, J.J.S. Oliveira JO83 (SP 445911!); São Paulo City, Parque Estadual das Fontes do Ipiranga, 1 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO329 (SP 445496!); ibid., 23 Mar. 2011, J.J.S. Oliveira & P.O. Ventura JO360 (SP 446016!); ibid., 2 Feb. 2012, J.J.S. Oliveira JO469 (SP 445554!); ibid., 18 Jan. 2013, N. Menolli-Jr. NMJ214 (SP 446032!); ibid., 14 Mar. 2014, M. Capelari (SP 446017!).

Notes — The material examined herein matches perfectly the description of the holotype (Wannathes et al. 2009). The record of M. suthepensis in both Thailand and Brazil may seem surprising, given the geographic distance and the presumed barriers to dispersal. However, a simple pairwise comparison of ITS sequences between the holotype from Thailand and the collections from Brazil showed almost no divergence (< 1 %) for this marker (Table S1), supporting the identification of the examined material.

It is possible that M. suthepensis have been introduced in South America from regions of Tropical Asia (or vice versa), supposing that the basidiospores or mycelium fragments were brought along with plants of economic interest historically exchanged between Asian and South American countries by merchant ships. Another hypothesis is the airborne spores and formation of spore clouds (Money 2016) that amplifies the power of spore discharge to cover long continental distances and transoceanic air trips. As fungal spores are constantly released to the air by many mushroom species, and although the vast majority are deposited in the surroundings, part of the discharged spores can be taken up by wind currents. However, spores of Marasmius are typically hyaline and thin-walled (Singer 1986, Desjardin 1989), highly susceptible to desiccation and UV radiation, and probably will not be viable after long aerial journeys (D.J. Lodge, pers. comm.).

Nonetheless, a thorough and specific biogeographic evaluation of pantropical distributions is necessary for many species exhibiting this supposedly wide distribution. As for M. suthepensis, other cases in Marasmius presenting supposedly wide distribution (showing molecular conspecificity) may suggest that the mycota of the Neotropics and Tropical Asia, which was connected in the remote past, still looks genetically linked. Many studies in Indo-Malaysian, Australasian, Oceanian and Afrotropical areas have already recorded species originally described in the Neotropics (Singer 1976, Corner 1996, Desjardin & Horak 1997, Desjardin et al. 2000, Antonín 2007, Tan et al. 2009, Wannathes et al. 2009, Antonín et al. 2012, Deng et al. 2012), but molecular comparison between strains indicating same morphological species is still in the beginning. We cannot discard the possibility that ITS is limited to solve species complex in allopatry, indicating that the samples represent very recently diverged taxa with little variation in particularly this marker.

Marasmius suthepensis is very close to M. neotropicalis in both morphology and genes herein studied. Moreover, M. suthepensis closely resembles M. ferruginoides (Antonín 2004), a species originally described from Tshopo Province (Democratic Republic of Congo) and also recorded from Cameroon, Ghana, Kenya, Nigeria (Antonín 2004, 2007). The examined material morphologically corresponds to M. ferruginoides according to Antonín (2004, 2007), diverging only by having slightly shorter pleurocystidia (21.3–36.3 vs 27–42 μm) while the holotype of M. suthepensis has pleurocystidia (27–47 μm) strongly concordant to those recovered from basidiomata of M. ferruginoides. The only difference is the number of lamellae: 12–18 with 3–4 lengths of lamellulae in M. suthepensis (holotype) vs 18–21 with 2–3 lengths of lamellulae in M. ferruginoides (holotype). The collections of M. suthepensis from Brazil show a wide variation in the number of lamellae (13–23), that overlaps both ranges above. Based on Shay et. al. (2017), M. ferruginoides (by collection from Madagascar) seems an independent species.

On the other hand, M. ferruginoides and M. suthepensis are micro-morphologically very similar to M. hinnuleus. Wannathes et al. (2009) argued that M. hinnuleus, described from Cuba, differs by having a sulcate to plicate, pale reddish brown to rufous cinnamon, smaller pileus (4–12 mm vs 10–27 mm) and longer basidiospores (mean 14 × 3.8 μm based on descriptions found in Singer (1976) and Pegler (1983)). Singer (1976) mentioned smaller basidiospores (7.7–)9.7–13.5 × 3.3–4.5 μm (generally 9.7–10.3 × 3.3–4.5 μm). The largest basidiospore length was given by Pegler (1983), 11–15.5 × 3.5–4.5 μm. The basidiospore dimensions in Singer and Pegler do not agree. Regarding the pileus colour, the protologue used the term ‘helvolus’, which is a variation of pale red to yellowish (Riddle 1870) and this pigmentation might have been observed in dried material but not in fresh material. Based on Shay et. al. (2017), M. hinnuleus (by collection from Madagascar) is an independent species.

TYS280 M. suthepensis (holotype) grouped along a basal tri- or polytomy with JO329 and JO469 in the /grammicola clade in all trees recovered. These three strains form a weakly supported branch because M. suthepensis seems to be part of a species complex that encompasses M. brunneoolivascens, M. conchiformis, M. griseoroseus, M. hobbitii, M. linderioides, M. luteoolivaceus and M. neotropicalis (Fig. 1b). Therefore, ITS alone is not sufficient to provide resolution for the clade, especially for M. suthepensis which is shown to be the earlier diverging lineage of the complex (see more in ‘Discussion’ section ‘Molecular phylogeny’).

Marasmius venatifolius J.S. Oliveira, sp. nov. — MycoBank MB811134; Fig. 17, ​,1818

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Fig. 17

Marasmius venatifolius (JO313). a. Basidiomata; b. basidiospores; c. basidioles; d. pleurocystidia; e. cheilocystidia; f. Siccus-type broom cells of the pileipellis. — Scale bars: a = 16.4 mm, b–f = 10 μm.

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Fig. 18

Marasmius venatifolius (JO313). Branched structures or broom cells of the stipitipellis. — Scale bars: 10 μm.

Holotype. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, on rotten leaves and small twigs of dicotyledonous trees, 9 Oct. 2010, J.J.S. Oliveira & P.O. Ventura JO313 (SP 445485!).

Etymology. From the Latin venatis = veined and folius = lamellae, referring to the veined lamellae.

Pileus 17.8–32 mm diam, sulcate-striate or irregularly sulcate, rufous orange or bright orange. Lamellae free or subfree, regular to intervenose. Stipe central, cylindrical, with a strigose basal mycelium. Basidiospores 6–9 × 3.5–5.3 μm, ellipsoid to subellipsoid. Pleurocystidia 27–69 × 5–12 μm, elongate, versiform, deeply immersed in the trama. Hyphae trama dextrinoid. Pileipellis hymeniform, with Siccus-type broom cells. Stipitipellis covered with forked to multibranched segments.

Pileus 17.8–32 mm diam, often convex-applanate, sometimes campanulate, sulcate-striate or slightly sulcate or even slightly tessellated, centre flat, shallowly depressed or slightly umbonate, margin decurved, plane to uplifted, edge regular, wavy or lacerate; centre dark orangish brown (Y₉₉M₆₀C₄₀) to purplish brown (N₇₀Y₇₀M₇₀) or almost black, becoming ochraceous to rufous orange (Y₉₉M₆₀C₃₀, N₃₀Y₉₉M₆₀ or N₄₀Y₉₉M₇₀), or bright orange (Y₉₉M₆₀C₂₀), to paler orange (Y₉₉M₅₀C₃₀) at the margin; membranous, context thin (< 1 mm); surface glabrous, barely pruinose, dry, dull, subvelutinous, non-hygrophanous. Lamellae free or subfree, subdistant to distant, L = 10–15, unequal, regular (only in JO285, but this clearly has some slightly veined lamellae) or mostly strongly interveined or veined, l = 1–2, opaque, smooth, pale cream (N₀₀Y₁₀M₀₀), edge even, partially or finely concolorous with the pileus as well as with the hymenium between the lamellae. Stipe 19–72 × 0.7–1.7 mm, central, cylindrical, equal, with a cream to orangish, strigose basal mycelium; tough, semicartilaginous to chitinous; apex concolorous with the lamellar faces, varying from yellowish brown (N₃₀Y₁₀M₃₀), to orangish brown (N₄₀Y₆₀M₅₀ to N₆₀Y₉₉M₇₀), reaching dark brown or black at the base; surface glabrous, finely pruinose near the base, smooth, opaque or silky bright.

Basidiospores (5.3–)6–9(–10) × 3.5–5.3(–7) μm (x_rm = 7–8 × 4.3–4.5 μm, x_mm = 7.6 (± 0.7) × 4.4 (± 0.1) μm, Q_rm = 1.7–1.8, Q_mm = 1.8 (± 0.1), n/s = 30, s = 2), obovoid, ellipsoid to subellipsoid, lacrimoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 23–36 × 4–8 μm, clavate, rarely tapering to the apex or somewhat mucronate, some apparently having a granular coating on the apex, hyaline, thin-walled, inamyloid. Pleurocystidia 27–69 × (3.4–)5–12 μm, elongate, versiform, clavate, vermiform, filamentous, some capitate or mucronate, generally opaque, often deeply rooted in the lamellar trama, projecting a little above the basidia and basidioles, smooth, inamyloid, sometimes inconspicuous. Cheilocystidia similar to the Siccus-type broom cells of the pileipellis, pale orange when grouped, main body 10–18.8 × 5.3–8.8(–13.4) μm, clavate to turbinate, sometimes forked, walls somewhat thinner; setulae apical, erect, generally elongate, 4.6–10.6 × 0.6–1.3(–2) μm, filiform, very thin to broad, regular in outline, solid, apex acute or needle-like. Lamellar trama dextrinoid, especially in the inner layer, irregular, hyphae interwoven, cylindrical, 2–14.4 μm diam, regular in outline, branched, smooth, hyaline, thin-walled. Pileus trama dextrinoid, especially in the subhymenium, irregular, hyphae interwoven, cylindrical, 2–10 μm diam, regular in outline, branched, smooth, hyaline, thin-walled. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, pale orange when grouped, hyaline when isolated, main body (8.8–)11.3–25 × 6.3–11.3(–14.3) μm, clavate to often turbinate, sometimes branched, rarely irregular in outline, thick-walled at the apex, narrowed downward to the base; setulae apical, erect, short to elongate, 2.5–8.3 × 0.6–1.3 μm, cylindrical, filiform, very thin, needle-like, simple, pale yellow, apex acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, 3–15 μm diam, regular in outline, yellowish brown or pale golden brown, smooth, thick-walled; internal hyphae hyaline, thin-walled, 2–10 μm diam. Stipitipellis covered by forked or multibranched segments, dendroid, present especially at the basal region of the stipe, irregular in outline, sometimes as a different type of broom cells, ramifications clavate or similar to digitiform diverticula, horn-shaped, cylindrical, rising from segments or hyphal endings, thick-walled, relatively short, 8–28.8 × 2.5–8.8 μm, tending to disappear or diminish upward to the stipe apex. Clamp connections present in almost all tissues, some small and inconspicuous, but not seen in the cortical stipe trama.

Habit & Substrate — Gymnopoid, scattered or gregarious, on rotten leaves or small twigs of dicotyledonous trees.

DNA Barcode — GenBank KP635203 (ITS1-5.8S-ITS2) and KP635157 (LSU).

Additional specimens (paratypes) examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 18 Mar. 2010, J.J.S. Oliveira JO63 (SP 445417!); ibid., 7 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO285 (SP 445467!).

Notes — Marasmius venatifolius shares some morphological similarities with M. spissus (Desjardin 1989), especially the pruinose stipe, abundant basal mycelium, the shape and length of basidiospores (5.6–8 μm), inconspicuous pleurocystidia, and the dendroid elements in the stipitipellis. Otherwise, they are very distinct species. Marasmius sullivantii is somewhat similar to the new species, especially in the dimension and pigmentation of the pileus, in the general aspect of the stipe, the shape and length of the basidiospores and the presence of broom cells in the stipitipellis (Desjardin 1989). However, M. venatifolius has a brighter orange, not ‘soredia-like’ spotted pileus, which is slightly tessellated due to the presence of intervenose lamellae; broader basidiospores (3.5–5.3(–7) μm vs 3.2–4 μm); differently shaped, deeper ‘rooted’ pleurocystidia; and more irregular broom cells bearing more elongate setulae, not accompanied by smooth cells. Marasmius venatifolius is a distinct species, arising as a well-supported branch of a polytomy in the Dataset 2.2 tree (Fig. 1c) and in the supported branch with an unresolved placement in the Dataset 3 tree (Fig. 2).

4 — Marasmius sect. Globulares Kühner emend. Antonín & Noordel. group Sicci Singer ser. Leonini Singer

Marasmius ambicellularis J.S. Oliveira, sp. nov. — MycoBank MB800886; Fig. 19

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Fig. 19

Marasmius ambicellularis (JO144). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. cheilocystidia: e1. Siccus-type broom cells, e2. smooth cells; f. pileipellis: f1. Siccus-type broom cells, f2. smooth cells.— Scale bars: a = 2.8 mm, b–f = 10 μm.

Holotype. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, on hanging dead palm leaves, 23 May 2010, J.J.S. Oliveira & C.L.A. Pires JO144 (SP 417465!).

Etymology. Based on the two types of cells in the pileipellis.

Differs from Marasmius linderioides by the absence of pleurocystidia, the presence of both smooth and Siccus-type broom cells in the pileipellis and the substrate type which is on hanging dead palm leaves.

Pileus 1–3 mm diam, convex, shallowly sulcate, centre plane, margin incurved to decurved, edge regular; pale orange (N₀₀Y₅₀M₂₀), with centre brownish orange (N₁₀Y₅₀M₆₀); membranous, context thin (< 1 mm); surface glabrous, dry, dull, non-hygrophanous. Lamellae adnate, distant, L = 8–10, equal, l = 1, faces and edges pale cream (N₀₀Y₁₀M₀₀). Stipe 1–2.5 × 0.3–0.5 mm, eccentric or central, reduced, curved, cylindrical, thin, chitinous, hollow; apex concolorous with the lamellae, middle orangish brown (N₂₀Y₈₀M₆₀), with a darker base; surface glabrous, smooth, opaque; basal mycelium scarce, cream, strigose.

Basidiospores 12.3–17 × 3.8–5 μm (x_m = 14.2 (± 1.2) × 4 (± 0.3) μm; Q_m = 3.5 (± 0.4), n = 30), suboblong, lacrimoid or subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia 18.8–31 × 7.3–8.8 μm, clavate, hyaline, thin-walled, 4 sterigmata, inamyloid. Basidioles 21.3–31.3 × 6–8 μm, clavate or acuminate, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia inconspicuous, similar to the elements of the pileipellis, dimorphic: 1) Siccus-type broom cells, main body 11.3–18 × 6.3–8 μm, clavate to vesiculose, or even irregular in outline, hyaline, thin-walled, inamyloid; setulae apical, erect, generally short, 0.9–4.6 × 0.6–1.6 μm, cylindrical or conical, sometimes rare, regular in outline or a little monilioid, hyaline, simple or branched, solid, apex obtuse and rounded; 2) Globulares-type smooth cells, 16.3–24 × 6.5–9 μm, very similar to the cells of the pileipellis. Lamellar trama dextrinoid, irregular, composed of cylindrical hyphae, 1.6–9.4 μm diam, regular in outline, smooth, hyaline, thin-walled, branched. Pileus trama similar to the lamellar trama, hyphae 1.6–10 μm diam. Pileipellis hymeniform, composed of dimorphic elements: 1) Siccus-type broom cells, abundant, main body 9.4–20 × 6.6–15 μm, clavate, vesiculose or turbinate, occasionally bilobed or irregular in outline, hyaline, thin- to moderately thick-walled, inamyloid; setulae apical, erect, short to a little elongate, 1.3–6.5 × 0.9–1.6 μm, cylindrical or conical, regular in outline to almost moniliform, branched, pale yellow to brown, solid, apex obtuse and rounded; 2) Globulares-type smooth cells, 15–24 × 8.5–14.3 μm, pyriform, less numerous, hyaline, thin- to moderately thick-walled. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, 3–7.5 μm diam, smooth, regular in outline, branched, melleous, thick-walled; internal hyphae similar to the cortical hyphae, but more slender, 1.5–6 μm diam, irregular, hyaline, thin-walled. Clamp connections present in all tissues.

Habit & Substrate — Marasmioid, gregarious, on hanging dead palm leaves.

DNA Barcode — GenBank KP635181 (ITS1-5.8S-ITS2) and KP635136 (LSU).

Notes — Marasmius ambicellularis is a distinct species by not having morphological resemblance to any currently known species, except for M. linderioides (Oliveira et al. 2014). However, M. linderioides differs by having larger pileus (up to 8 mm diam); by more numerous lamellae (12–15) and 2–3 series of lamellulae; by shorter basidiospores (10–15 μm long); by the presence of well-developed pleurocystidia; and by having only Siccus-type broom cells in the pileipellis; with basidiomata generally found on substrate on the ground. Additionally, M. ambicellularis was collected in the end of the rainy season equivalent to late fall while M. linderioides was found in early summer (the beginning of the rainy season) suggesting they might differ seasonally. Also distinguishing, M. linderioides was found producing more gregarious and abundant basidiomata on the substrate, suggesting somewhat different spore dispersal strategy. On the other hand, both species are sympatric, collected near each other in the same area and occupying similar niches, decomposing dried palm debris.

No significant phylogenetic distinction was found between the collections of M. ambicellularis and M. linderioides in all phylogenetic trees (Fig. 1b, ​,2).2). A divergence of 0.2 % between ITS sequences of the respective collections may reveal that ITS is perhaps not sufficiently distinct to grant genetic differentiation of these two species. New data (more collections and more genetic markers) regarding this species complex are needed to clarify the molecular systematics of these taxa.

Marasmius bellus Berk., Hooker’s J. Bot. Kew Gard. Misc. 8: 139. 1856 — Fig. 20

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Fig. 20

Marasmius bellus (JO299). a. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. Siccus-type broom cells of the pileipellis. — Scale bars: a = 22.2 mm, b–e = 10 μm.

≡ Chamaeceras bellus (Berk.) Kuntze, Revis. Gen. Pl. (Leipzig) 3: 455. 1898.

Pileus 16–28 mm diam, convex to applanate, striate or slightly to strongly sulcate-striate, centre flat, margin decurved to straight, edge crenate to wavy; whitish cream to yellowish cream (N₀₀Y_10–30M₀₀), with a pale brown (N₁₀Y₆₀M₃₀) small spot at the centre; membranous, context thin (< 1 mm); surface glabrous, dry, dull, papyraceous, non-hygrophanous. Lamellae free to adnexed, subclose to subdistant, L = 14–18, equal, regular or rarely with few veins, l = 3, opaque, smooth, cream (N₀₀Y₁₀M₀₀), edge even, concolorous with the lamellar faces. Stipe 25–44 × 0.8–1.2 mm, central, cylindrical to almost filiform, equal, tough, chitinous, hollow; apex concolorous with the lamellar faces, latter orangish brown (N₄₀Y₇₀M₆₀) at the middle of the stipe, becoming black toward the base; surface glabrous, smooth, opaque; basal mycelium abundant, pale cream, strigose.

Basidiospores 8–13.3 × 3.3–4.5 μm (x_m = 9.8 (± 0.8) × 3.8 (± 0.3) μm, Q_m = 2.6 (± 0.3), n = 30), oblong, clavate, subfusoid to lacrimoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 20–27 × 4.8–7 μm, clavate to rarely subacute at the apex, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia present on the sterile lamellar edge, hyaline, Siccus-type broom cells, hyaline to pale yellow, main body 11.3–19 × 5–8(–12.5) μm, clavate to slightly turbinate, hyaline, thin-walled; setulae apical, erect, generally elongate, sometimes short, 2–7 × 0.5–1 μm, cylindrical, filiform, needle-like, solid, simple, hyaline, apex acute or rarely obtuse. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 1.6–12 μm diam, regular in outline, branched, smooth, hyaline, thin-walled. Pileus trama dextrinoid, irregular, hyphae similar to those of the lamellar trama, 2–12.5 μm diam. Pileipellis hymeniform, dextrinoid, Siccus-type broom cells, abundant, orange or ochraceous in mass, completely hyaline when isolated, difficult to separate, delicate, easily breakable at the base, main body (6.3–)10–18 × 6.3–14.4 μm, clavate or more often turbinate, sometimes flattened, rarely branched, hyaline, thin-walled; setulae apical, erect, short to elongate, 2–10.3 × 0.4–1.3 μm, filiform, setiform or needle-like, thin, regular in outline, solid, sometimes with lumen, simple, hyaline to pale yellow, apex acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, 2.5–11.3 μm diam, regular in outline, smooth, melleous, darkening to olivaceous yellow or almost black, thick-walled; internal hyphae hyaline, cylindrical, 2.5–9.4 μm diam, smooth, thin-walled. Clamp connections present in almost all tissues except the cortical stipe trama.

Habit & Substrate — Gymnopoid to marasmioid, gregarious, on all sorts of debris of dicotyledonous tree.

Distribution — The species was originally described from Amazonas State (Brazil), and it has also been recorded from Pernambuco (Singer 1976) and São Paulo (Pegler 1997) States. Other records of the species are from Bolivia (Singer 1976), Venezuela (Dennis 1961), Trinidad, Martinique, Dominique (Pegler 1983) and Puerto Rico (Lodge 2003, Cantrell et al. 2006).

Specimen examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 8 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO299 (SP 445478!).

Notes — This is the second record of M. bellus from Reserva Biológica de Paranapiacaba. The material herein examined is concordant with descriptions provided by Dennis (1951a), Singer (1976) and Pegler (1983, 1997). The description in Singer (1976) was based on the holotype (also in Singer 1958, 1965), accompanied by specimens more recently collected from Brazil and Bolivia. However, Singer (1976) described the specimens as having a little larger pileus (13–35 mm diam), adnate lamellae (in some cases), and presence of dissimilar basidioles. Dennis (1951a) provided a revised description of the type and reported adnate and slightly interconnected lamellae. He also identified specimens from Venezuela (Dennis 1961) as M. bellus, reporting the presence of poorly developed pleurocystidia and apparently shorter setulae on broom cells of the pileipellis. The presence of pleurocystidia is, though, conflicting with all descriptions found in Singer and Pegler studies, and even in the revision of the holotype accomplished by Dennis (1951a).

Pegler (1983) mentioned neither pleurocystidia nor basidioles and described the lamellae as being adnexed to adnate, moderately sized pileus (35 mm diam), and basidiospores ranging from 11–14 μm long. Finally, the characteristics mentioned by Pegler (1997) from the material collected from Campinas City and Santo André City (São Paulo State) agree with the material herein examined. Marasmius cremeus (which is almost identical to M. similis) is a similar species, but differs by the more delicate and smaller pileus (3–14 mm diam), thinner stipe (0.2–0.5 mm) and by the presence of larger basidiospores ((11–)12–14(–16) μm) (Wannathes et al. 2009).

In the Dataset 3 trees (Fig. 2), M. bellus has an unsupported relationship with M. leoninus, M. ruber and M. trinitatis. Marasmius bellus could be confused with pale pigmented specimens of M. leoninus, but does not have a distinctive taste whereas M. leoninus tastes very disagreeable (D.J. Lodge, pers. comm).

Marasmius cladophyllus var. glaberripes Singer, Fl. Neotrop. Monogr. 17: 214. 1976 — Fig. 21

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Fig. 21

Marasmius cladophyllus var. glaberripes (JO87). a. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. Siccus-type broom cells of the pileipellis. — Scale bars: a = 16.6 mm, b–e = 10 μm.

Pileus 9–55 mm diam, globose when young, becoming hemispheric, convex to plane at maturity, sometimes becoming revolute, generally smooth, sometimes slightly tessellated (reticulate-striate), faintly rugose at the centre or with margin slightly sulcate-striate, centre flat to slightly depressed, margin incurved to straight, or somewhat uplifted, edge regular; dark orange when young (N₂₀Y₉₉M₉₀), becoming orange (N₀₀Y₉₉M₄₀), yellowish orange (N₁₀Y₉₉M₆₀) to darker orange (N₁₀Y₉₉M₉₀) or reddish orange (N₄₀Y₉₉M₆₀), centre dark ochraceous (N₂₀Y₉₉M₅₀) to even darker (near N₄₀Y₉₉M₆₀); membranous, context white to brownish cream, thin to thickish (≤ 1 mm); surface glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae sinuate to adnate, close to subdistant, L = 12–14, unequal, strongly reticulate to merulioid-anastomosed, becoming intervenose near the pileus edge, l = 3–4, opaque, smooth, pale cream (N₀₀Y₁₀M₀₀), pale yellow (N₀₀Y₃₀M₀₀) to pale beige (N₀₀Y₂₀M₁₀), edge even, concolorous or finely discolorous with the lamellar faces (with a thin orange line), hymenium between lamellae cream. Stipe 38–72 × 1–3 mm, central, cylindrical, equal or with a little flared apex, sometimes compressed; tough, chitinous to more semicartilaginous, hollow; apex concolorous with the lamellar faces, sometimes cream 2/3 over the basal 1/3 of the stipe, becoming orange or ochraceous (N₆₀Y₉₉M₇₀) or brown (N₂₀Y₉₀M₆₀), reaching dark reddish brown (N_60–80Y_90–99M_70–80) downward to the base; surface glabrous, slightly pruinose at the base, smooth, opaque or silky bright; basal mycelium strigose, abundant, white to sordid yellow.

Basidiospores 8.8–11.3 × 2.5–3.8 μm (x_m = 10 (± 0.8) × 3 (± 0.3) μm; Q_m = 3.2 (± 0.4), n = 30), oblong, clavate to subfusoid, hyaline, thin-walled, smooth, inamyloid. Basidia not observed. Basidioles 17.5–28.8 × 4–7 μm, clavate, sometimes with apex tending to acuminate, hyaline, thin-walled. Pleurocystidia absent. Cheilocystidia abundant and conspicuous, in form of Globulares-type smooth cells, 22.5–32 × 8–10 μm, pyriform or vesiculose, hyaline, thin-walled. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2–7.5 μm diam, smooth, hyaline, thin-walled, septate. Pileus trama hyphae similar to those of the lamellar trama, hyphae cylindrical, 2.3–11.3 μm diam. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, pale orange when in group, main body 7.5–23.8 × 5–8.8 μm, broadly cylindrical, clavate to turbinate, almost pedicellate, rarely lobed or branched, hyaline, thin-walled; setulae apical, erect, 3.8–10 × 0.6–1.3 μm, cylindrical, filiform, thin, regular in outline, simple, pale orange, abundant, apex acute. Stipe trama dextrinoid, cortical hyphae parallel, 7–15 μm diam, regular in outline, cylindrical, smooth, pale yellow to pale brown, thick-walled, septate; internal hyphae hyaline, 3.8–6.3 μm diam, thin-walled, slender. Clamp connections present in all tissues.

Habit & Substrate — Gymnopoid, gregarious or scattered, on leaves and branches of dicotyledonous trees.

Distribution — This variety of M. cladophyllus was originally described from Tungurahua, Ecuador (Singer 1976). This is the first record from Brazil.

Specimens examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 16 Dec. 2009, M. Capelari & L.A.S. Ramos 4562 (SP 445784!); ibid., L.A.S. Ramos & M. Capelari 86 (SP 445424!); ibid., 24 Mar. 2010, J.J.S. Oliveira JO87 (SP 445424!); Iporanga City, Parque Estadual Turístico do Alto Ribeira, Núcleo Santana, 29 Feb. 2012, J.J.S. Oliveira & D.E. Desjardin JO518 (SP 445587!); ibid., D.E. Desjardin DED8681 (SP 445664!).

Notes — Marasmius cladophyllus is easily recognized in the field due to its typical merulioid-anastomosed lamellae or almost favoloid hymenophore. The species currently contains four varieties: cladophyllus, glaberripes, intermedius and tjibodensis. Marasmius cladophyllus var. glaberripes differs from var. cladophyllus by having completely glabrous stipe and, therefore, by the absence of caulocystidia, also by having shorter basidiospores (barely reaching 10 μm long) and cheilocystidia in form of dextrinoid hyphal ends that are conspicuously contorted and often forked (Singer 1976). Besides the presence of hyaline, thin-walled, versiform or contorted, often cylindrical or ventricose, some diverticulate, hair-like caulocystidia (35–50 × 4–9.8 μm) accompanied by ventricose cells with or without a mucro, var. cladophyllus also presents, in contrast to var. glaberripes, somewhat longer basidiospores (up to 12.7 μm) and intermittent clusters of regular broom cells interspersed with versiform or hyphal ends at the lamellar edge (Singer 1976). Mainly because of the glabrescent stipe, the collections examined are determined as belonging to var. glaberripes, in spite of having a little longer basidiospores and cheilocystidia only in the form of Globulares-type cells.

On the other hand, M. cladophyllus var. intermedius (Singer 1976) has a subglabrous stipe, beset by minute hyphoid excrescences that are occasionally similar to hyaline setulae and few broader elements like those found in the type variety. Additionally, it has cheilocystidia of two types: a) fusoid and long beaked (23–26 × 5–6 μm); and b) vesiculose, often mucronate, hyaline, broadened (12–27 × 6–18 μm) and inamyloid; and dimorphic trama: a) strongly dextrinoid, with thick-walled hyphae; or b) inamyloid to weakly dextrinoid, with thin-walled hyphae. Marasmius cladophyllus var. tjibodensis, a fourth variety, was described by Desjardin et al. (2000) and it is similar to var. glaberripes in the type of cheilocystidia. But var. tjibodensis is distinguished by having a pruinose and fibrillose stipe and, thus, it has caulocystidia as well as longer basidiospores (up to 13.5 μm).

The presence of elongate caulocystidia suggests that M. cladophyllus var. cladophyllus is a member of series Atrorubentes in the traditional view. Singer (1976) formally classified the species within ser. Actinopodes, later separated into ser. Atrorubentes and ser. Spinulosi (currently morphological concepts). But Singer’s decision remained inconclusive since he stated that, because of the varieties glaberripes and intermedius, the species would belong to series Leonini (Singer 1976). This morphological incongruence is problematic since var. glaberripes cannot be classified in ser. Atrorubentes at all, but would be properly placed in ser. Leonini.

In the trees (Fig. 1b, ​,2),2), M. cladophylus var. glaberripes is an independent basal branch of a polytomy, near but poorly related to other taxa including M. leoninus, thus supporting Singer’s (1976) alternative suggested placement of M. cladophyllus in ser. Leonini. Moreover, MCA1837 M. cf. cladophyllus, also included in the analyses of Group 1, is a genetically distinct species. ITS sequences retrieved from GenBank and identified as M. cladophyllus were not included in the analyses because of their poor quality (JF767003, KF241549, HQ248212, AY216475, HQ248211).

Marasmius corrugatus (Pat.) Sacc., Syll. Fung. (Abellini) 16: 54. 1902 — Fig. 22

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Fig. 22

Marasmius corrugatus (JO466). a. Basidiomata; b. basidiospores; c. basidia; d. basidioles; e. cheilocystidia; f. pileipellis: f1. small, more numerous, thin-walled Siccus-type broom cells; f2. larger, less numerous, thick-walled Siccus-type broom cells; g. structures of the stipitipellis. — Scale bars: a = 19.7 mm, b–g = 10 μm.

≡ Androsaceus corrugatus Pat., Bull. Soc. Mycol. France 16: 175. 1900.

≡ Marasmius corrugatus (Pat.) P. Syd., Just’s Bot. Jahresber. 28: 132. 1902.

Pileus 3.2–28 mm diam, hemispheric when young, convex to more often plane, sometimes revolute, orbicular or sometimes almost lobed, centre flat or slightly depressed, sometimes shallowly umbonate, smooth or radially corrugated, margin non-striate or slightly sulcate-striate, decurved, plane or slightly uplifted, edge regular or sometimes lacerate; brownish orange (N₅₀Y₉₉M₅₀) when young, becoming yellowish brown or pale orangish brown (N₃₀Y₈₀M₄₀), yellowish orange (N₂₀Y₇₀M₄₀) or pale orange (N₃₀Y₆₀M₃₀) to orange (N₂₀Y₈₀M₅₀), centre ochraceous or orangish brown (N₃₀Y₉₀M₅₀ to N₄₀Y₉₉M_50–60), sometimes almost cream (N₀₀Y₅₀M₁₀ to N₁₀Y₅₀M₂₀) toward the margin; membranous, context cream, thin (< 1 mm); surface glabrous, dry, sometimes a little humid, dull, subvelutinous, rarely micaceous, non-hygrophanous. Lamellae free, adnexed or adnate, close to subdistant, abundant, L = 28–36, unequal, regular, some forked, l = 4–6, opaque, smooth, whitish cream (N₀₀Y₁₀M₀₀) to pale yellow (N₂₀Y₇₀M₄₀), edge even, concolorous with the lamellar faces. Stipe 6–55 × 0.5–3 mm, central, cylindrical, equal or slightly broader at the apex or base, sometimes compressed, tough, semicartilaginous to chitinous, hollow; apex concolorous with the lamellae, becoming pale brownish orange (N_20–30Y₇₀M₄₀) or even darker (N₄₀Y_80–99M₅₀ to N₅₀Y₉₀M₇₀) at the middle, reaching to dark reddish brown (N₈₀Y₉₉M₇₀ to N₉₀Y₉₉M₇₀) or black at the base; surface glabrous, smooth, shiny with a silky sheen; basal mycelium whitish cream to orangish, strigose or tomentose.

Basidiospores 6.3–11.3(–12.5) × 3–5.3(–6) μm (x_rm = 8.3–9.5 × 3.7–4.6 μm; x_mm = 8.8 (± 0.4) × 4 (± 0.4) μm, Q_rm = 1.8–2.6, Q_mm = 2.2 (± 0.3); n/s = 30, s = 5), obovoid to more often suboblong, lacrimoid, subellipsoid, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia 17.5–26.3 × 4.4–7.5 μm, clavate, smooth, hyaline, 2–4 sterigmata, thin-walled, inamyloid. Basidioles 16.6–23.8 × 4.4–7.5 μm, clavate to subfusoid, smooth, hyaline, thin-walled. Pleurocystidia absent. Cheilocystidia in form of Siccus-type broom cells, completely hyaline, inconspicuous and delicate in some specimens, otherwise conspicuous, but not numerous; main body 10–25 × 4.8–9.4 μm, cylindrical-clavate, clavate to slightly turbinate, some forked, hyaline, thin- to slightly thick-walled, inamyloid; setulae apical, erect, short to elongate, 3.8–15 × 0.8–2 μm, cylindrical, filiform or flagellum-like, simple, sometimes contorted, solid, hyaline, apex acute. Lamellar trama strongly dextrinoid becoming vinaceous to ferruginous in Melzer’s reagent, some with an inner trama more strongly dextrinoid, completely disorganized or irregular, hyphae interwoven, cylindrical, 2.5–12.5 μm diam, regular in outline, some segments inflated, smooth, branched, hyaline, thin-walled. Pileus trama strongly dextrinoid, tending to have a regular arrangement (parallel), slightly interwoven in some places, trama completely irregular in some specimens, hyphae cylindrical, 2–12.5 μm diam, regular in outline, some segments a little inflated. Pileipellis hymeniform, orange or ochraceous, mottled, composed of Siccus-type broom cells in two types: 1) abundant, smaller in proportion, pale orange or brownish ochre when in group, main body 12.5–17.5(–20) × 5.6–12 μm, cylindrical-clavate, clavate to turbinate, rarely forked, thin-walled, hyaline, dextrinoid; setulae apical, erect, short to elongate, 1.3–16.3 × 0.5–2.5 μm, cylindrical, filiform, needle-like, or digitiform, regular in outline, simple to rarely branched, hyaline to pale orange, solid or with lumen, apex acute to slightly obtuse; 2) larger broom cells, less numerous in proportion, but regularly distributed, more irregular in outline, main body 12–26.5 × 5.6–12.5 μm, clavate to slightly turbinate, thick-walled, melleous, apex pale orange, hyaline at the base; setulae apical, erect, short and often very elongate, 3.8–35 × 1.3–5 μm, cylindrical, digitiform, needle-like, simple to branched, solid or with lumen, melleous or pale orange, apex obtuse or acute. Stipe trama dextrinoid, cortical hyphae parallel, dark brown to black, becoming olivaceous green, orange or even melleous yellow when isolated, cylindrical, regular in outline, 2.5–13.8 μm diam, smooth, thick-walled; internal hyphae cylindrical, 1.3–11.3 μm diam, disorganized, hyaline, thin-walled, with numerous septa. Stipitipellis without broom cells, but with short structures, 10–23.8 × 2–7 μm, clavate to conical, thick-walled, seemingly hyphal endings, or lateral sprouts from cortical hyphae, apex slightly acuminate, but generally obtuse. Clamp connections present in almost all tissues, but not seen at the base of the second broom cells type of the pileipellis.

Habit & Substrate — Gymnopoid, gregarious, sometimes almost caespitose, growing on twigs and leaves of dicotyledonous trees.

Distribution — Originally described from Guadeloupe, it is also known from Venezuela, Bolivia, Argentina (Singer 1976), Martinique, Dominique (Pegler 1983), Puerto Rico (Lodge 2003, Cantrell et al. 2006) and Brazil (Pegler 1997).

Specimens examined. Brazil, São Paulo City, Santo André City, Reserva Biológica de Paranapiacaba, 24 Mar. 2010, J.J.S. Oliveira JO85 (SP 445423!); São Paulo City, Parque Estadual das Fontes do Ipiranga, 1 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO336 (SP 445500!); ibid., 3 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO347 (SP 445507!); ibid., 2 Feb. 2012, J.J.S. Oliveira JO466 (SP 445551!); Parque Estadual da Cantareira, Núcleo Engordador, 30 Jan. 2012, J.J.S. Oliveira & V. Motato-Vásquez JO456 (SP 445543!).

Notes — In addition to the type variety of M. corrugatus, three other varieties were established by Singer (1976): a) var. aurantiacus which differs by forming smaller basidiomata, more orangish pileus, narrower basidiospores (3–3.7 μm broad) and by having well-developed broom cells in the stipitipellis; b) var. lacustris, similar in pileus pigmentations to var. aurantiacus, but diverging from all by forming much larger pileus (up to 57 mm diam) and by having dimorphic cystidia in the stipitipellis; and c) var. portonovensis which differs from the type variety by having more smooth cells than broom cells in the pileipellis and on the lamellar edge, by having strongly dimorphic lamellar trama and shorter basidiospores (6.2–7.5 μm). Desjardin & Horak (1997) established M. corrugatus var. olivaceobrunneus from New Guinea, which differs by having olivaceous brown pileus and pruinose stipe, covered by caulocystidia in form of broom cells.

The specimens analysed correspond to the type variety as described by Singer (1976), differing only by the smaller pileus (up to 28 mm diam) and by the absence of smooth cells in the pileipellis. The presence of smooth cells mixed with broom cells, in some species, may be considered an occasional characteristic, since it can be present or absent depending on the population or on the stage of development of the basidiomata. The mottled aspect of the pileipellis is due to a gradual pattern of broom cells from small, less pigmented and thin-walled to large, more pigmented and thick-walled. Pegler (1983) also studied the holotype along with collections from Martinique and Dominique, finding larger basidiospores (9–13 × 2.7–4 μm). Marasmius maculosus (Takahashi 2002) is very similar to M. corrugatus, differing only by the reddish brown pileus and stipitipellis with clusters of thin hyphae.

Marasmius corrugatus grouped within a strongly supported, well-defined subclade (Fig. 1b, ​,2)2) named /corrugatus, which also includes M. congregatus and M. pseudoniveoaffinis.

Marasmius dimorphus C. Puccin. & Capelari, Mycotaxon 95: 298. 2006 — Fig. 23

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Fig. 23

Marasmius dimorphus (JO298). a. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. pileipellis: e1. regular Siccus-type broom cells, e2. irregular, thick-walled Siccus-type broom cells. — Scale bars: a = 17.6 mm, b–e = 10 μm.

Pileus 4–37 mm diam, hemispheric, conical or campanulate, but generally convex, especially when mature, sulcate, centre flat or slightly umbonate, margin decurved, edge regular to crenate; light cinnamon (N₃₀Y_50–60M₃₀) to ochraceous brown (N₄₀Y₆₀M₄₀ to N₅₀Y₇₀M₄₀) or pale pinkish brown or almost beige (N₂₀Y_40–50M₃₀), generally light brown (N₃₀Y₄₀M₂₀), some pale hazel brown (N₆₀Y_70–80M₅₀) or cocoa (N₇₀Y_50–60M₄₀), becoming darker greyish brown (N₆₀Y₄₀M₃₀) when dried, with a dark brown centre (N₉₀Y₉₉M_60–80, N₉₉Y₉₉M₇₀ or Y₉₉M₆₀C₅₀) or paler (N₉₀Y₉₉M₆₀); membranous, context thin (< 1 mm); surface glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae free to adnexed, subclose to distant, L = 14–18, equal, regular, rarely forked, thin, l = 0–1, opaque, smooth, cream (N₀₀Y₁₀M₀₀) to pale beige (N₀₀Y₃₀M₁₀), edge even, concolorous with the lamellar faces, hymenium between the lamellae partially concolorous with the pileus. Stipe 18–80 × 0.4–1 mm, central, cylindrical, sometimes thin, equal, tough, hardly chitinous, hollow; apex concolorous with the lamellae, becoming brownish beige (N₁₀Y₄₀M₂₀) or golden brown (N₄₀Y₅₀M₄₀), reaching to dark brown (N₉₀Y₉₉M_50–60) or to black toward the base; surface glabrous, smooth, glossy; basal mycelium abundant, light cream, strigose or tomentose.

Basidiospores 11.3–17.5(–18.8) × 3–5 μm (x_rm = 13.9–15.4 × 3.8–4.5 μm; x_mm = 14.7 (± 0.6) × 4.2 (± 0.3) μm, Q_rm = 3.7–4.5, Q_mm = 4.1 (± 0.4); n/s = 30, s = 4), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 18.8–27.5 × 5–8 μm, clavate, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia in form of Siccus-type broom cells, faintly brown to hyaline in mass, main body 8.8–15 × 3.8–7.5 μm, clavate to slightly turbinate, regular in outline, apex with thick walls, but thin-walled at the base, breaking easily under gentle pressure on the coverslip, hyaline, inamyloid; setulae apical, erect, 3–8 × 0.8–1 μm, digitiform or cylindrical, regular or somewhat irregular in outline, with ramifications, pale brown to hyaline, solid, apex obtuse to acute. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2.5–11.3 μm diam, regular in outline, smooth, hyaline, some segments slightly inflated, thin-walled. Pileus trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2.5–11.3 μm diam, similar to those of the lamellar trama. Pileipellis hymeniform, mottled, composed of Siccus-type broom cells in two types: 1) abundant, brownish when in group, main body 5–18.8 × 5–8(–11.3) μm, clavate to turbinate, sometimes flattened, or with an elongate base, brown, apex with thick wall, but thinning toward the base; setulae apical, erect, 4.4–7.5 × 0.8–1.3 μm, digitiform to cylindrical, filiform, regular in outline, pale brown, simple, solid, apex obtuse to acute; 2) irregular in outline, rare, completely brown or dark melleous, main body 12.5–38.8 × 5–9.8 μm, irregular in outline, tending to clavate, with thicker walls; setulae apical, erect, 5–10 × 1.3–2.5 μm, more elongate and broader compared with type 1, solid, apex obtuse. Stipitipellis dextrinoid, cortical hyphae parallel, cylindrical, 3–13.8 μm diam, regular in outline, smooth, light brown to melleous, walls moderately thick, septa numerous and close; internal hyphae hyaline, 2–11.3 μm diam, thin-walled, smooth. Clamp connections present in almost all tissues, inconspicuous in the pileus trama and not seen in the lamellar trama.

Habit & Substrate — Marasmioid, gregarious, on twigs, thin branches and dried leaves of dicotyledonous trees.

Distribution — The species is known only from São Paulo State (Puccinelli & Capelari 2006).

Specimens examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 8 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO298 (SP 445477!); São Paulo City, Parque Estadual das Fontes do Ipiranga, 1 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO334 (SP 445499!); ibid., 3 Mar. 2011, J.J.S. Oliveira & F. Karstedt JO355 (SP 445513!); Parque Estadual da Cantareira, Núcleo Engordador, 19 Dec. 2011, J.J.S. Oliveira & M. Capelari JO442 (SP 445535!); ibid., 30 Jan. 2012, J.J.S. Oliveira & V. Motato-Vásquez JO462 (SP 445548!).

Notes — Marasmius dimorphus was described from Parque Estadual das Fontes do Ipiranga, and it also occurs in Reserva Biológica de Paranapiacaba and Parque Estadual da Cantareira, areas that are currently devoid of forest connections. The material examined is completely compatible with the original description in Puccinelli & Capelari (2006).

Marasmius helveloides (Singer 1976) is a similar species with a pale brown pileus and lacking pleurocystidia, but differs by having more robust stipe (1–2 mm thick) that is indistinctly pruinose due to the presence of broom cells in the stipitipellis; by the absence of dimorphic brooms cells in the pileipellis; and by the presence of rare cystidioles.

Marasmius dimorphus grouped within the strongly supported subclade /rhabarbarinus, along with M. corneri, M. rhabarbarinoides and M. rhabarbarinus (Fig. 1c). These species are also morphologically close and may represent a species complex (Greenwood 1974). The phylogenetic position is kept in the Dataset 3 tree (Fig. 2), with even higher support because M. coneri is absent.

Marasmius graminicola Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 6: 111. 1899 — Fig. 24

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Fig. 24

Marasmius graminicola (JO459). a1, a2. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. Siccus-type broom cells of the pileipellis. — Scale bars: a1 = 10 mm, a2 = 5 mm, b–e = 10 μm.

Pileus 3–7.5 mm diam, conical-campanulate to parabolic-convex, smooth to striate or shallowly sulcate-striate, more strongly sulcate-striate when mature or dried, centre slightly to strongly umbonate, margin decurved, edge regular; pale orange (N₀₀Y₅₀M₂₀) to pale ochraceous (N₁₀Y₇₀M₃₀), orange (N₁₀Y₉₀M₄₀) to brownish orange (N₃₀Y₉₉M₅₀), with centre intensely orange (N₄₀Y₉₉M₆₀) to dark brownish orange (N₅₀Y₉₉M₆₀) or concolorous with the margin; membranous, context cream, thin (< 1 mm); surface glabrous, dry, dull, papyraceous to subvelutinous, non-hygrophanous. Lamellae free to subfree, or almost adnexed, subclose to subdistant, L = 13–16(–20), equal, regular, l = 1–2, opaque, smooth, whitish cream (N₀₀Y₁₀M₀₀), edge even, faintly marginate and finely brownish orange or not marginate and concolorous with the lamellar face, as well as with the hymenium between the lamellae. Stipe 33–60 × 0.2–1 mm, central, cylindrical, sometimes filiform, equal, semi cartilaginous to chitinous, hollow; apex pale yellow, becoming brownish orange (N₅₀Y₉₀M₅₀), reaching dark amber brown (N₅₀Y₉₉M₅₀) or dark brown (N₈₀Y_90–99M₅₀) below; surface glabrous, smooth, with a silky sheen; basal mycelium yellowish, strigose.

Basidiospores 10–14.4 × 3–4.4 μm (x_rm = 11.8–12.6 × 3.6–4 μm; x_mm = 12.2 (± 0.6) × 3.8 (± 0.3) μm, Q_rm = 3–3.5, Q_mm = 3.3 (± 0.4); n/s = 30, s = 2), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 22.5–28.8 × 5–7.5 μm, cylindrical, clavate to often fusoid, often with an acute apex, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia in form of Siccus-type broom cells similar to those of the pileipellis, cinnamon in mass, abundant, main body 10–22.5 × (3.8–)5–7.5 μm, cylindrical to clavate, sometimes a little turbinate, thin-walled; setulae apical, erect, 3–7.5 × 0.8–1.3 μm, conspicuous, cylindrical or digitiform, some very elongate, setoid, regular in outline, simple, pale yellow, solid, apex acute. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 1.6–12.5 μm diam, regular or irregular in outline, sometimes inflated, branched, smooth, thin-walled. Pileus trama similar to the lamellar trama, hyphae cylindrical, 2–15 μm diam. Pileipellis hymeniform, composed of Siccus-type broom cells, pale yellow when in group, main body 8.8–22 × 5.6–15 μm, clavate or more frequently turbinate, sometimes ventricose or bilobed, hyaline, thin-walled; setulae apical, erect, 2.5–8 × 0.6–1 μm, filiform, setiform, cylindrical, or digitiform, simple, regular in outline, hyaline, apex generally acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, regular in outline, 2.5–7.5 μm diam, smooth, yellowish melleous in KOH, thick-walled; internal hyphae with similar dimensions, also parallel, hyaline, smooth, thin-walled. Clamp connections present in almost all tissues, absent in the cortical stipe trama.

Habit & Substrate — Marasmioid, gregarious, on leaf petioles and dried twigs of dicotyledonous trees.

Distribution — The species was originally described from Buenos Aires, Argentina, and it was also reported from Mexico (Singer 1976) and Republic of South Korea (Antonín et al. 2012). This is the first record of the species from Brazil.

Specimens examined. Brazil, São Paulo State, São Paulo City, Parque Estadual da Cantareira, Núcleo Engordador, 30 Jan. 2012, J.J.S. Oliveira & V. Motato-Vásquez JO459 (SP 445546!); ibid., 9 Feb. 2012, J.J.S. Oliveira & M. Capelari JO480 (SP 445559!); Parque Estadual das Fontes do Ipiranga, 2 Feb. 2012, J.J.S. Oliveira JO473 (SP 445916!).

Notes — The collections examined diverge from the description of the species provided by Singer (1976) by having a more elongate stipe (up to 60 vs up to 40 mm) and lamellae more numerous (13–16 vs 8–9), also presenting lamellulae, which are typically absent. Moreover, no dark orange or reddish brown pileus was observed in the examined material and the lamellar edge tends to be only faintly marginate. Singer (1976) also noted that the species was always associated with monocotyledonous plants, mostly Gramineae, but spreading to other nearby material.

Antonín et al. (2012) recorded the species from South Korea, describing the collections with stipes much longer (20–100 mm), basidiospores more elongate (12–15(–17.5) μm), lamellae more numerous ((10–)16–18(–24)) and growing only on dicotyledonous leaves. Marasmius bambusiniformis is another similar species, but differs by having larger basidiospores ranging from 15–18.5 × 3.7–5 μm (Singer 1976, Desjardin & Horak 1997, Tan et al. 2009, Wannathes et al. 2009).

A pairwise comparison of the ITS region (Table S2), including sequences of M. graminicola obtained by Antonín et al. (2012), M. bambusiniformis obtained by Wannathes et al. (2009), and sequences generated from the examined material revealed that the samples of M. graminicola collected from São Paulo are not conspecific with the samples of M. graminicola in Antonín et al. (2012), so reports from Korea are likely a misapplied name. Sequences by Antonín et al. (2012) of M. ‘graminicola’ from Korea were nearly identical with those of M. bambusiniformis in Wannathes et al. (2009). Marasmius graminicola grouped within /graminicola subclade (Fig. 2) as sister to ‘M. luteoolivaceus + M. hobbitii’ branch with a high support (PP 0.99).

Marasmius hobbitii J.S. Oliveira, sp. nov. — MycoBank MB811135; Fig. 25

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Fig. 25

Marasmius hobbitii (JO309). a. Basidiomata; b. basidiospores; c. basidia, d. basidioles; e. cystidioles; f. cheilocystidia; g. broom cells of the pileipellis. — Scale bars: a = 11.7 mm, b–g = 10 μm.

Holotype. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, on fallen palm leaves, 9 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO309 (SP 445484!).

Etymology. This species is named after the Hobbits, from the book ‘The Hobbit’ by J.R.R. Tolkien, a race of usually mycophile, very tiny people (c. 60–120 cm tall), passing unseen by common human eyes and with hairy feet. The small size and hairy feet of the hobbits connect them to the reduced size of the basidiomata and the strigose mycelium (hair) at the base (foot) of the stipes found in M. hobbitii.

Pileus 1.8–4 mm diam, campanulate, slightly sulcate, orange. Lamellae free, subdistant (7), equal, non-marginate. Stipe 13–15 × 0.3–0.5 mm, central, curved, filiform, with a strigose basal mycelium. Basidiospores 13.8–19(–20) × 5–9 μm, fusoid-ventricose, rhomboid. Basidioles clavate, along with acuminate cystidioles. Pleurocystidia absent. Pileipellis made up by Siccus-type boom cells, with apical setulae 1.5–4.6 × 0.5–1 μm.

Pileus 1.8–4 mm diam, campanulate, sometimes hemispheric, shallowly sulcate, centre flat, margin decurved, edge regular or crenate; orange (N₂₀Y₉₉M₅₀) to paler orange (N₁₀Y₉₉M₄₀) toward the margin, centre somewhat darker orange or rufous (N₃₀Y₉₀M₆₀); membranous, context thin (< 1 mm); surface glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae free, subdistant, L = 7, equal, regular, l = 0, opaque, smooth, pale cream (N₀₀Y₁₀M₀₀), edge even, concolorous with the lamellar faces as well as with the interlamellar hymenium. Stipe 13–15 × 0.3–0.5 mm, central, curved or oblique, thin, filiform, equal; pliant, chitinous, hollow; apex concolorous with the lamellar face, otherwise dark brown all over; surface glabrous, smooth, opaque; basal mycelium cream-coloured, strigose.

Basidiospores 13.8–19(–20) × 5–9 μm (x_rm = 17–18 × 6.6–7.3 μm; x_mm = 17.5 (± 0.7) × 7 (± 0.5) μm, Q_rm = 2.5–2.6, Q_mm = 2.6 (± 0.1); n/s = 30, s = 2), oblong, fusoid-ventricose, rhomboid, distinctly broad, smooth, hyaline, thin-walled, inamyloid. Basidia 25–36.3 × 8.8–12 μm, clavate, smooth, hyaline, thin-walled, 2–4 sterigmata, inamyloid. Basidioles 25–35 × 5.6–12 μm, clavate or broadly clavate, along with acuminate cystidioles, with a tapered, or even with a rostrate and acute apex, orthochromatic in Crezil blue like the basidia and basidioles, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia in form of Siccus-type broom cells, very similar to those in the pileipellis, abundant, also occasionally found dispersed throughout the lamellar faces, hyaline, main body 11.3–21.3 × 5.5–10 μm, clavate to slightly turbinate, thin-walled; setulae apical, erect, short, 1–3.8 × 0.8–1 μm, digitiform to verruciform, solid, simple, hyaline, apex obtuse to slightly acute. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2–8.8 μm diam, regular in outline, branched, smooth, hyaline, thin-walled. Pileus trama similar to the lamellar trama, hyphae 2.5–12.5 μm diam. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, orange or pale ochraceous in mass, hyaline when isolated, main body 10–20 × 7–13.8 μm, clavate to slightly turbinate, rarely ventricose, sometimes branched, hyaline, thin-walled, dextrinoid; setulae apical, erect, generally short, some a little more elongate, 1.5–4.6 × 0.5–1 μm, verruciform to digitiform, simple or branched, hyaline, solid, apex acute to slightly obtuse. Stipe trama weakly dextrinoid, but the reaction is noticeable only in the internal hyphae, cortical hyphae parallel, cylindrical, 2.5–7.5 μm diam, regular in outline, smooth, septate, melleous or brown in KOH, thick-walled; internal hyphae parallel or disorganized, 1.3–6.3 μm diam, thin-walled. Clamp connections present in almost all tissues, inconspicuous, not observed in the cortical stipe trama.

Habit & Substrate — Marasmioid, gregarious on fallen palm leaves.

DNA Barcode — GenBank KP635180 (ITS1-5.8S-ITS2) and KP635135 (LSU).

Additional specimen (paratypes) examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 8 Jan. 1991, L.K. Okino & M. Capelari 10 (SP 445761!).

Notes — Marasmius hobbitii is characterized by small basidiomata with an orange, often campanulate pileus, by thin, short stipes, by broad basidiospores, 5–9 μm in width, and by the presence of very acuminate cystidioles. It is a little difficult to define whether the cystidioles may or may not be considered pleurocystidia in KOH, which would put the species in sect. Sicci ser. Haematocephali according to Singer (1976). When the lamellae sections are mounted in Cresyl Blue, all cells present in the hymenial layers are orthochromatic (non-reactive or blue), indicating that the longer, acuminate cells should be cystidioles indeed. Marasmius hobbitii shows close relationship (PP 1.0/BS 89) with M. luteoolivaceus in the Dataset 3 tree (Fig. 2), within the /graminicola subclade.

Marasmius persicinus (Desjardin et al. 2000) is morphologically similar, differing by having a much paler pileus, more numerous lamellae (10–12), slender basidiospores (3–4 μm in width), and by the absence of acuminate cystidioles. Marasmius hobbitii is also comparable with M. bambusiniformis (Singer 1976) and M. pusio (Singer 1976). However, M. bambusiniformis, while also occurring on monocotyledonous plants, has a more ferruginous, larger pileus (4.5–7 mm diam), a more thickened stipe, a pubescent stipe base, and narrower basidiospores (3.7–5 μm). Based on Singer (1976), M. pusio differs by having more numerous lamellae (8–12) which may be regularly didymous or tridymous (forked twice or three times respectively), by the pallid or paler stipe, by the differently shaped, smaller basidiospores (6–10.3 × 2.5–4.5(–5.5) μm), by the pileipellis with branching Ramaria-like broom cells (illustrated and defined in Singer (1965, 1976)), many of them pedicellate, and by the stipitipellis containing broom cells. Despite the differences noted above, both M. bambusiniformis and M. pusio have hymenial cystidioles resembling those in M. hobbitii.

Marasmius leoninus Berk., Hooker’s J. Bot. Kew Gard. Misc. 8: 135. 1856 — Fig. 26

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Fig. 26

Marasmius leoninus (JO84). a. Basidiomata: a1. top view of the pileus, a2. whole basidiomata, a3. hymenophore view; b. basidiospores; c. basidioles; d. cheilocystidia; e. Siccus-type broom cells of the pileipellis. — Scale bars: a1, a3 = 9.3 mm, a2 = 16.7 mm, b–e = 10 μm.

Pileus 9–22 mm diam, convex to plane, or almost revolute, tending to form lobes, smooth or slightly sulcate-striate at the margin, centre slightly depressed, margin decurved to straight, edge regular or slightly wavy; orange (N₃₀Y₉₀M₅₀) to pale orange (N₀₀Y₇₀M₅₀ to N₂₀Y₆₀M₄₀), brownish orange (N₅₀Y₉₉M₇₀) or sometimes with diffuse, brownish (N₃₀Y₈₀M₆₀) spots or shades at the centre; membranous, but sometimes apparently cartilaginous or coriaceous, context white, thin (< 1 mm); surface glabrous, dry, dull, subvelutinous, non-hygrophanous. Lamellae free to adnexed, subclose, L = 20–22, equal, l = 2–3, some forked, opaque, smooth, pale cream (N₀₀Y₃₀M₀₀), edge even, concolorous with the lamellar faces or finely concolorous with the pileus. Stipe 40–57 × 1–1.5 mm, central, cylindrical, sometimes compressed, slightly flared at the apex, contorted; cartilaginous to chitinous, tough, hollow; apex concolorous with lamellar faces, becoming brownish orange (N₄₀Y₉₀M₅₀) or brown (N₃₀Y₉₀M₇₀), reaching to dark reddish brown (N_80–90Y₉₉M₇₀) or almost black at the base; surface glabrous, but finely pruinose under lens, opaque; basal mycelium densely strigose, with yellow or orange tints.

Basidiospores 7–10(–11.3) × 2.5–4.6 μm (x_rm = 8.7–9.3 × 3.6–4.1 μm; x_mm = 9 (± 0.4) × 3.9 (± 0.4) μm, Q_rm = 2.1–2.7, Q_mm = 2.4 (± 0.4); n/s = 30 from JO84 and 20 from JO320, s = 2), obovoid to suboblong, ellipsoid, lacrimoid or subclavate, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 21.3–32 × 3.8–6.3 μm, cylindrical, clavate, some subacuminate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia in form of Siccus-type broom cells, somewhat similar to those of the pileipellis, completely hyaline, main body 13.3–22.5 × 5–8.8 μm, clavate to slightly ventricose, thin- to thick-walled; setulae apical, erect, longer and broader than those on the pileipellis broom cells, 2.5–12.5 × 0.6–1.5 μm, cylindrical, filiform, setoid, regular in outline, hyaline, solid, simple, apex acute. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2.5–13.8 μm diam, regular or irregular in outline, some inflated, branched, smooth, thin- to thick-walled. Pileus trama strongly dextrinoid, irregular, hyphae interwoven, cylindrical, 1.6–15 μm diam, often irregular in outline, inflated, branched, heteromorphic, some thick- and other thin-walled, smooth. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, pale orange in mass, becoming hyaline when isolated, main body 12.5–25 × 5.6–17.5 μm, clavate to turbinate, sometimes ventricose, or subglobose, hyaline; setulae apical, erect, 2–6.3 × 0.5–1.3 μm, abundant, cylindrical, filiform, digitiform or setiform, orange, simple, regular in outline, hyaline or pale yellow, solid, apex acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, regular in outline, 2.5–11.3 μm diam, smooth, sometimes branched, brown, thick-walled; internal hyphae hyaline, 3–7.5 μm diam, thin-walled. Stipitipellis with structures that resemble Siccus-type broom cells, with setulae digitiform to cylindrical. Clamp connections present in all tissues.

Habit & Substrate — Gymnopoid, gregarious, on rotten twigs of dicotyledonous trees.

Distribution — The species was originally described from Amazonas State, Brazil, and it has also been recorded from the United States, Venezuela, Ecuador, Bolivia, Argentina (Singer 1965, 1976), Martinique, Dominique, Guadalupe, Trinidad (Pegler 1983) and Puerto Rico (Lodge 2003, Cantrell et al. 2006). In Brazil, the species is also known from Rio Grande do Sul (Singer 1976) and São Paulo States (Pegler 1997).

Specimens examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 24 Mar. 2010, J.J.S. Oliveira JO84 (SP 445422!); ibid., 9 Dec. 2010, J.J.S. Oliveira, P.O. Ventura & A.V. Costa JO320 (SP 445489!).

Notes — Marasmius leoninus has a wide range of variation in morphology. For example, according to Singer (1976), the species varies in pruinosity of the stipe, arrangement of the pileus trama and in the presence or absence of cheilocystidia, and in many other characteristics such as colour, shape, size and texture of the pileus, and the pattern of lamellae attachment to the apex of stipe. D.J. Lodge (pers. comm.) has consistently observed strongly disagreeable taste in all her collections of M. leoninus.

The examined material fits within Singer’s (1965, 1976) concept in almost all characteristics, only differing by having somewhat smooth to slightly sulcate-striate pileus margin instead of distinctly sulcate and by having more numerous lamellae. The pruinosity of the stipe surface due to the presence of broom cells and the dimorphic pileus trama make JO84 and JO320 very compatible with the species. However, neither of these are very strong characteristics for separation of taxa (D.J. Lodge, pers. comm.).

Marasmius leoninus var. aberrans differs from the material examined by having a foliicolous habit, completely glabrous stipe and slightly shorter basi-diospores (up to 9.5 μm). Singer (1976) synonymized M. orinocensis (≡ Androsaceus orinocensis) under M. leoninus, but he noticed that it might be considered a variety (M. leoninus var. orinocensis ad interim) combining the presence of dimorphic pileus trama (thin-walled and thick-walled hyphae) and growth on wood (lignicolous) in contrast to monomorphic trama (thin-walled hyphae) and foliicolous (growing on leaves) in var. leoninus. According to the above-mentioned characteristics, the specimens herein examined would fit within the var. orinocensis concept. However, the formal separation into var. leoninus and var. orinocensis was not accomplished. Except for a possible substrate preference, this distinction based on the characteristics of the pileus trama seems very inconclusive.

Pegler (1983) recorded specimens of M. leoninus having apricot yellow to cinnamon, glabrous, radially sulcate-striate pileus (10–30 mm diam), glabrous and glossy stipe, subfusoid basidiospores (8–10 μm long), presence of cheilocystidia and absence of broom cells in the stipitipellis, growing on leaves. He also reported the slightly thickened wall for the hyphae of the context (pileus trama). Pegler (1983) also supported the identification of the species by arguing that the number of lamellae is a variable feature in M. leoninus and corresponds to the protologue (Berkeley 1856).

Marasmius leoninus seems to be closer to M. bellus and M. ruber, but without a defined resolution or significant support values on the phylogenetic trees of Datasets 2.1 and 3 (Fig. 1b, ​,22).

Marasmius rhabarbarinoides J.S. Oliveira, sp. nov. — MycoBank MB811136; Fig. 27

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Fig. 27

Marasmius rhabarbarinoides (JO66). a. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. pileipellis: e1. irregular Siccus-type broom cells, e2. ordinary Siccus-type broom cells. — Scale bars: a = 14.6 mm, b–e = 10 μm.

Holotype. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, on dead leaves of dicotyledonous trees, 20 Mar. 2010, J.J.S. Oliveira JO66 (SP 445418!).

Etymology. The name refers to Marasmius rhabarbarinus, based on the resemblance to that species.

Pileus 23 mm diam, hemispheric, shallowly sulcate, centre dark brown or orange brown, then ochraceous or carrot orange. Lamellae free, subdistant (15), pale cream. Stipe 63 × 1 mm, central, cylindrical, dark brown, with a scarce, tomentose basal mycelium. Basidiospores 20–25.4 × 3.3–4(–4.8) μm, subclavate or fusoid. Pleurocystidia absent. Hyphal trama strongly dextrinoid. Pileipellis hymeniform, mottled, composed of thin- or thick-walled Siccus-type broom cells.

Pileus 23 mm diam, convex or almost hemispheric, shallowly sulcate, centre wrinkled, slightly umbonate, margin incurved, edge regular; orangish brown (N₂₀Y₉₉M₆₀), reaching to bright ‘carrot’ orange (N₁₀Y₉₉M₇₀) with a dark brown centre (N₆₀Y₉₉M₈₀); membranous, context thin (< 1 mm); surface glabrous, dry, dull, somewhat micaceous, velutinous, non-hygrophanous. Lamellae free, subdistant, L = 15, equal, l = 0, smooth, pale cream (N₀₀Y₂₀M₀₀), edge partially concolorous with the pileus, but also with the lamellar faces. Stipe 63 × 1 mm, central, cylindrical, equal, tough, chitinous, hollow; apex concolorous with the lamellar face, going to dark brown or black elsewhere down to the base; surface glabrous, smooth, glossy; basal mycelium scarce, cream-coloured, tomentose.

Basidiospores 20–25.4 × 3.3–4(–4.8) μm (x_m = 22.7 (± 1.3) × 3.8 (± 0.3) μm; Q_m = 6 (± 0.5), n = 30), oblong, subclavate or often fusoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 23–34.8 × 5–8.3 μm, cylindrical to clavate, some with a tapered apex, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia similar to type 1 of the Siccus-type broom cells of the pileipellis, but main body proportionally smaller, abundant, cream when in group, 13.5–21.3(–25) × 6–8.8(–10.3) μm, clavate to turbinate, sometimes ventricose, hyaline, thin-walled; setulae apical, erect, abundant, 3–6 × 0.8–1.5 μm, cylindrical, regular or irregular in outline, solid or with lumen, apex acute or slightly obtuse. Lamellar trama strongly dextrinoid, irregular or disorganized, hyphae interwoven, cylindrical, 1.8–12.5 μm diam, regular in outline, branched, hyaline, thin-walled. Pileus trama similar to the lamellar trama, hyphae cylindrical, 2.3–17 μm diam. Pileipellis hymeniform, mottled, composed of Siccus-type broom cells, abundant, consisting of two types: 1) less numerous and more conspicuous, main body more elongate, (13.8–)17.5–30 × 5.3–8.8(–11) μm, subcylindrical, clavate, a little turbinate, sometimes irregular in outline, thin-walled at the base, but strongly thick-walled at the middle toward the apex, golden yellow to orangish yellow, colourless at the base; setulae apical, erect, 4.3–10 × 0.8–1.6 μm, digitiform to cylindrical, regular in outline, some branched, solid, pale golden yellow, apex acute or somewhat obtuse; 2) more numerous and less conspicuous, yellowish when in group, main body smaller, 8.8–17.5 × 5.3–13.8 μm, clavate to turbinate, sometimes bilobed, or irregular in outline, thin-walled, hyaline; setulae apical, erect, 2.8–9 × 0.8–1.3 μm, digitiform to cylindrical, thin, simple or branched, regular in outline, solid, sometimes with lumen, tapering upwards, apex acute or somewhat obtuse. Stipe trama strongly dextrinoid at least at the apex, cortical and internal hyphae similar, cylindrical, 2–18.8 μm diam, regular in outline, branched, smooth, light chestnut brown, walls not very thick or almost thin, septa numerous. Clamp connections present in all tissues.

Habit & Substrate — Marasmioid, solitary, on dead leaves of dicotyledonous trees.

DNA Barcode — GenBank KP635190 (ITS1-5.8S-ITS2) and KP635145 (LSU).

Notes — Marasmius rhabarbarinoides differs from M. rhabarbarinus mainly by having longer basidiospores (20–25.4 vs 14.4–21.5 μm). Although morphologically similar, M. rhabarbarinoides is 4.9–5.9 % divergent from representatives of M. rhabarbarinus based on data ITS (Table S3), supporting the establishment of the new species. In all phylogenetic trees (Fig. 1, ​,2),2), M. rhabarbarinoides is sister to M. rhabarbarinus, with a high support within the subclade /rhabarbarinus.

Marasmius rhabarbarinus Berk., Hooker’s J. Bot. Kew Gard. Misc. 8: 135. 1856 — Fig. 28

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Fig. 28

Marasmius rhabarbarinus (JO494). a. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. Siccus-type broom cells of the pileipellis. — Scale bars: a = 21.4 mm, b–e = 10 μm.

≡ Chamaeceras rhabarberinus (Berk.) Kuntze, Revis. Gen. Pl. (Leipzig) 3: 457. 1898.

Pileus 1.3–45 mm diam, hemispheric or slightly conical, tending to campanulate, orbicular, shallowly to deeply sulcate, centre flat to slightly umbonate, margin incurved to decurved, edge regular to crenate; ochraceous (N₃₀Y₉₉M₅₀), ferruginous orange (N₂₀Y₉₉M₇₀), ‘carrot’ orange or ‘rhubarb’ fulvous orange (N₄₀Y₉₀M₆₀ to N₄₀Y₉₉M₆₀), or brownish orange (N₅₀Y₉₉M₆₀), with a dark orangish brown or brown centre (N_60–90Y₉₉M₇₀ to N₉₀Y₉₉M_70–80); membranous, context pale cream, thin (< 1 mm); surface glabrous, dry, dull, rarely discreetly semitranslucent, papyraceous to subvelutinous, non-hygrophanous. Lamellae free, subclose to distant, L = 12–20, equal, regular, seldom slightly intervenose, l = 1–3, opaque, smooth, pale cream (N₀₀Y₁₀M₀₀) to pale yellow (Y₄₀M₁₀C₀₀), edge even, concolorous with the lamellar face or very finely brownish orange, hymenium between lamellae partially concolorous with the pileus. Stipe 35–98 × 1–3 mm, central, cylindrical, sometimes compressed, base slightly broader, tough, semicartilaginous to chitinous, hollow; apex concolorous with the lamellar faces or a little bit more yellowish (N₁₀Y_60–70M₂₀, N₀₀Y₈₀M₂₀), reaching to pale brown (N₄₀Y₇₀M₄₀) or darker brown (N₆₀Y₉₉M₆₀) and blackish brown (N₉₀Y₉₉M_60–70) to black toward the base; surface glabrous, smooth, having a silky shine, with a slightly pubescent region restricted to the base; basal mycelium greyish cream, tomentose.

Basidiospores (12–)14.4–21.5 × 3–4.8(–6.8) μm (x_rm = 18–20 × 3.8–4 μm, x_mm = 18.5 (± 0.7) × 3.9 (± 0.1) μm, Q_rm = 4.7–5, Q_mm = 4.9 (± 0.1), n/s = 30, s = 5), oblong, clavate to subfusoid, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 20–28 × 5–7.3 μm, clavate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia in form of Siccus-type broom cells, hyaline, inconspicuous, main body 8.8–20 × 4.4–9.5(–11.3) μm, clavate, subcylindrical, turbinate, rarely pyriform, hyaline, thin-walled; setulae apical, erect, short to elongate, 1–7.5 × 0.6–1 μm, cylindrical, digitiform or filiform, hyaline, solid, simple, apex obtuse or acute. Lamellar trama distinctly dextrinoid, but inamyloid in the subhymenium, irregular, hyphae interwoven, cylindrical, 1.3–14.4 μm diam, regular in outline, some segments inflated, smooth, hyaline, thin-walled. Pileus trama dextrinoid, irregular to regular, or organized over the subhymenium, hyphae cylindrical, 2.5–13.8 μm diam, other characteristics similar to those of the lamellar trama. Pileipellis hymeniform, mottled, composed of Siccus-type broom cells, abundant, pale yellowish orange when in group, especially due to the setulae; main body (7–)10–25.6 × 5.3–11.3 μm, clavate to pyriform, sometimes flattened, short to elongate, generally regular in outline: 1) apex solid with a densely thick wall, golden yellow or yellowish, wall thinner toward the base; 2) hyaline and thin-walled all over; setulae apical, erect, short to very elongate, 4.6–15 × 0.6–1.6(–2) μm, regular in outline, cylindrical, digitiform to filiform, some conical, melleous or yellowish, hyaline and thinner on those entirely thin-walled, apex obtuse or acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, 5–12.5 μm diam, regular in outline, smooth, not branched, ochre or orangish brown, some segments paler and somewhat olivaceous, or very pale to completely hyaline at the stipe apex, moderately thick-walled; internal hyphae hyaline, 3.8–12.5 μm diam, parallel. Clamp connections present in almost all tissues, absent in the cortical stipe trama.

Habit & Substrate — Marasmioid, gregarious, scattered or solitary, more frequently on the lamina or on the petioles of dead leaves, rarely on twigs of dicotyledonous trees.

Distribution — The species was originally described from Amazonas State (Brazil) and it is also known from São Paulo State (Singer 1976, Pegler 1997) and from Argentina (Singer 1976).

Specimens examined. Brazil, São Paulo State, São Paulo City, Parque Estadual da Cantareira, Núcleo Engordador, 5 Jan. 2012, M. Capelari & P.O. Ventura 4637 (SP 445783!); ibid., 30 Jan. 2012, J.J.S. Oliveira & V. Motato-Vásquez JO457 (SP 445544!); ibid., 9 Feb. 2012, J.J.S. Oliveira & M. Capelari JO474 (SP 445558!); ibid., J.J.S. Oliveira & M. Capelari JO475 (SP 445918!); ibid., J.J.S. Oliveira & M. Capelari JO476 (SP 445919!); ibid., 16 Feb. 2012, J.J.S. Oliveira & M. Capelari JO494 (SP 445565!).

Notes — The collections examined are not only concordant with the morphological description of the species by Singer (1976), but also add more information. The species has previously only been reported on dead leaves. Marasmius berteroi is similar, but has shorter spores (11–17 μm) and grows on mixtures of fallen leaves and wood (Singer 1976). Marasmius rhabarbarinus is also very close to M. rhabarbarinoides (previous taxon) and to M. dimorphus, this later differing by having pale brown to cocoa brown pileus, by having 0–1 series of lamellulae, and by having shorter basidiospores (11.3–17.5 (–18.8) μm). Moreover, M. rhabarbarinus is also difficult to distinguish from M. corneri when comparing their respective morphological descriptions. The unique (but not strictly) distinguishing characteristic is the absence of a mottled pileipellis in M. corneri, but the authors have mentioned the presence of thin- to thick-walled broom cells and setulae pale yellow to tawny, which may produce a mottle aspect (Wannathes et al. 2009). However, the pairwise comparison of ITS sequences (Table S3) confirms that these taxa are different species (c. 10 % divergence). This could be considered an example of cryptic species (Bickford et al. 2007), geographically and genetically separated, but morphologically similar.

Marasmius rhabarbarinus, along with M. rhabarbarinoides, M. corneri and M. dimorphus, comprise the highly supported subclade /rhabarbarinus (Fig. 1c, ​,22).

Marasmius ruber Singer, Sydowia 18: 342. 1965 — Fig. 29

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Fig. 29

Marasmius ruber (DED8669). a. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. Siccus-type broom cells of the pileipellis. — Scale bars: a = 13.6 mm, b–e = 10 μm.

Pileus 20 mm diam, convex to plano-convex or plane with a shallow central depression, striate, glabrous or subvelutinous, dull, dry, bright orange (6A7–8). Lamellae shallowly adnexed, subdistant with 3–4 series of lamellulae, narrow (< 2 mm), bright orange (6A7–8) with strongly reddish orange (7B8) edge. Stipe 35 × 1 mm, cylindrical, equal, fistulose, glabrous, when young light yellow (4A5–6) overall, in age apex light yellow, base brownish orange (6C5–6) to light brown (6D5–6); non-insititious, basal mycelium tawny strigose (macromorphological description and drawing (Fig. 29a) found in a small sheet along with the collection (unpublished), by Dennis E. Desjardin, 2012).

Basidiospores (6.3–)7.5–10.5 × 2–3.4 μm (x_m = 8.8 (± 1.1) × 3 (± 0.3) μm, Q_m = 3 (± 0.4), n = 30), oblong, slender, subfusoid or clavate, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 20–28 × 4.8–7 μm, narrowly cylindrical, clavate or acuminate, some with a pointed apex, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia in form of Siccus-type broom cells similar to those of the pileipellis, losing the reddish pigmentation in KOH, abundant, main body 15–18.8 × 5.6–8 μm, clavate to slightly turbinate, thin-walled; setulae apical, erect, 2–7.5 × 0.6–1 μm, cylindrical, filiform, regular in outline, hyaline, solid, simple, apex acute. Lamellar trama dextrinoid, irregular, hyphae interwoven, cylindrical, 2–7.5 μm diam, regular in outline, branched, smooth, thin-walled. Pileus trama dextrinoid, similar to the lamellar trama, hyphae 2–7.5 μm diam. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, distinctly orange to reddish when grouped, depigmenting in KOH, becoming hyaline, main body 12.5–22.5 × 3–12.5 μm, cylindrical, clavate to often turbinate, hyaline, thin-walled; setulae apical, erect, 2.5–7.5 × 0.5–1.3 μm, abundant, cylindrical, filiform, simple, regular in outline, solid, hyaline, apex acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, regular in outline, 3–15 μm diam, smooth, orangish, pale brown or melleous when depigmented in KOH, thick-walled; or a second type of worm-like hyphae present, rough or with shallow or small protuberances, or slightly incrusted, opaque, 2.3–8 μm diam, sometimes vein-like, conspicuous, with strongly thick walls, almost solid, aseptate, resembling skeletal hyphae, distinct within the trama; internal hyphae indistinguishable. Clamp connections present in almost all tissues, absent in the second type of hyphae in the cortical stipe trama.

Habit & Substrate — Gymnopoid, solitary, on dead twigs.

Distribution — Originally described from Beni (Bolivia), also known from Trinidad (Singer 1965, 1976) and from São Paulo State, Brazil (Grandi et al. 1984).

Specimen examined. Brazil, São Paulo, Iporanga City, Parque Estadual Turístico do Alto Ribeira, 28 Feb. 2012, leg. D.E. Desjardin DED8669 (SP 445657!).

Notes — The examined material corresponds to the description of the species by Singer (1965, 1976). An important observation is the presence of conspicuous, thick-walled hyphae, mixed within the stipe trama, which could correspond to the non-septate hyphae mentioned by Corner (1996) as skeletic in the stipe of M. dimiticus. Marasmius ruber appears in Group 1 (Fig. 1), and in an unsupported grouping formed by M. bellus, M. trinitatis and the /leoninus clade (Fig. 2) and its placement was not strongly supported in the trees recovered.

Marasmius trinitatis Dennis, Trans. Brit. Mycol. Soc. 34: 425. 1951 — Fig. 30

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Fig. 30

Marasmius trinitatis (JO306). a. Basidiomata; b. basidiospores; c. basidioles; d. cheilocystidia; e. Siccus-type broom cells of the pileipellis; f. Siccus-type broom cells of the stipitipellis. — Scale bars: a = 14.4 mm, b–f = 10 μm.

Pileus 6–34 mm diam, convex, tending to plane, smooth or slightly sulcate-striate, centre flat, margin incurved to plane, sometimes uplifted, edge regular or wavy; centre dark grey (N₈₀Y₃₀M₁₀) and margin paler grey (N₆₀Y₂₀M₁₀) when young, becoming olivaceous cinnamon (N₆₀Y₅₀M₂₀) or yellowish grey (N₅₀Y₇₀M₂₀) or brownish yellow (N₃₀Y₅₀M₂₀), with a greenish grey or brownish olive centre (N₇₀Y₅₀M₂₀) when mature; membranous, context thin (< 1 mm); surface glabrous, dry, dull to semitranslucent, subvelutinous, non-hygrophanous. Lamellae adnexed, adnate to sinuate, subdistant, L = 8–10, unequal, l = 3, opaque, smooth, light cream (N₀₀Y₁₀M₀₀), edge even, partially concolorous with the pileus, later becoming concolorous with the lamellar faces as well as with the hymenium between the lamellae. Stipe 8–40 × 0.9–1.5 mm, central, cylindrical, equal or a little broader at the apex, semicartilaginous to chitinous downwards, hollow; apex concolorous with the lamellar faces, becoming orangish brown (N₃₀Y₈₀M_50–60) at the middle, reaching dark brown (N₈₀Y₉₉M₆₀) at the base; surface glabrous, smooth, opaque; basal mycelium abundant, orangish yellow, strigose.

Basidiospores 10–13.3 × 3.4–4 μm (x_m = 11.7 (± 0.9) × 3.7 (± 0.2) μm; Q_m = 3.2 (± 0.3), n = 30), oblong, clavate, smooth, hyaline, thin-walled, inamyloid. Basidia not observed. Basidioles 23.8–30.6 × 4.5–7.5 μm, clavate, smooth, hyaline, thin-walled, inamyloid. Pleurocystidia absent. Cheilocystidia similar to the Siccus-type broom cell of the pileipellis, abundant, melleous in mass, main body 11.3–21.3 × 5–8.3 μm, cylindrical to clavate or turbinate, wall relatively thin, but thicker at the apex, hyaline; setulae apical, erect, short to elongate, 2.5–8 × 0.5–1.3 μm, cylindrical, filiform, needle-like, simple, solid, melleous to pale yellow, or hyaline, regular in outline, apex acute. Lamellar trama strongly dextrinoid, irregular, hyphae interwoven, cylindrical to inflated, slender to broad, 2.5–16.3 μm diam, branched, smooth, hyaline, thin-walled. Pileus trama strongly dextrinoid, irregular, hyphae interwoven, with some loose regions, cylindrical to inflated, 3.5–18.8 μm diam, like those of the lamellar trama. Pileipellis hymeniform, composed of Siccus-type broom cells, abundant, golden yellow to pale brown in mass, main body 9.4–19.4 × 5.6–10.6 μm, cylindrical to clavate or turbinate, walls relatively thin, but thicker above, melleous, or hyaline; setulae apical, erect, short to elongate, 1.5–7.5 × 0.4–1 μm, cylindrical, filiform, needle-like, simple, solid, melleous to pale yellow, or hyaline, regular in outline, apex acute. Stipe trama dextrinoid, cortical hyphae parallel, cylindrical, 3.5–11.3 μm diam, regular in outline, branched, smooth, dark melleous or brownish yellow, thick-walled; internal hyphae 2.5–20 μm diam, thin- to thick-walled, smooth. Stipitipellis containing Siccus- or Amiloflagellula-type broom cells, more frequent at the upper part of the stipe where the hyphae are more hyaline, other broom cells concolorous with the cortical hyphae of the median and basal part of the stipe, very variable in shape, setulae elongate in both types, apex acute. Clamp connections present in all tissues.

Habit & Substrate — Gymnopoid, gregarious, on rotten wood of dicotyledonous trees.

Distribution — Originally described from Trinidad (Dennis 1951c, Pegler 1983), it is also known from the United States, Mexico, Brazil, Bolivia (Singer 1965, 1976), Puerto Rico (Lodge 2003, Cantrell et al. 2006) and Papua New Guinea (Desjardin & Horak 1997). This is the first record of the species from São Paulo State.

Specimen examined. Brazil, São Paulo State, Santo André City, Reserva Biológica de Paranapiacaba, 9 Dec. 2010, J.J.S. Oliveira & P.O. Ventura JO306 (SP 445481!).

Notes — The examined material corresponds to M. trinitatis as described by Singer (1976), differing only by the occasional presence of pale melleous cheilocystidia, and the presence of broom cells in the stipitipellis. The latter characteristic is found in M. trinitatis var. immarginatus, but this variety does not possess cheilocystidia (Singer 1976). Perhaps the presence of broom cells in the stipitipellis is an occasional characteristic in this species and/or the absence of cheilocystidia in var. immarginatus might have been due to the condition of the material examined by Singer (1976). Because of this conflict between the presence of cheilocystidia and the main distinguishing characteristic of var. immarginatus (absence of cheilocystidia), the specimen is herein determined as the type variety. Regarding the pigmentation of the cheilocystidia, it was observed that the lamellar edge is concolorous with the pileus in the young basidiomata, but fading at the maturity, when the cheilocystidia become hyaline.

Marasmius trinitatis is basal to M. coarctatus and M. cladophyllus in the group 1 tree of Dataset 2.1 (Fig. 1b), but with weak statistical support. In the Dataset 3 tree (Fig. 2), M. trinitatis is basal to M. leoninus in /leoninus, but with weak support. In both analyses, the clade with M. trinitatis is placed in a polytomy.

Morphology and infrasectional inconsistences

In the traditional view, sect. Sicci sensu Singer is composed of species having the pileipellis composed of Siccus-type broom cells, in rare cases mixed with Globulares-type smooth cells. Within sect. Sicci Singer, series Atrorubentes is defined by the presence of caulocystidia and the absence of setae (Desjardin & Horak 1997), series Spinulosi is composed of species with setae in the pileus, lamellae and/or stipe (Antonín & Noordeloos 1993), species in series Haematocephali share the presence of pleurocystidia and absence of setae and caulocystidia (but may have broom cells in the stipitipellis) (Singer 1976), and series Leonini is characterized by the absence of all those features mentioned above, with the exception of broom cells in the stipitipellis (Singer 1976). Section Globulares sensu Singer is composed of species having the pileipellis solely composed of Globulares-type smooth cells. However, some special cases of known species or new morphological data recovered from recently described species bring inconsistences in those concepts, especially from tropical areas where diversity of species and morphology is high and still mostly unknown.

The presence of elongate caulocystidia as determinant for sect. Sicci ser. Atrorubentes may be remarkable in some species (i.e., M. atrorubens), but inexpressive in others (i.e., M. congregatus). Regarding the inexpressive forms of caulocystidia, it is also hard to handle with the reduced structures in the stipitipellis in M. corrugatus, classified within sect. Sicci ser. Leonini. Another example is the case of M. cladophyllus var. glaberripes (with a completely glabrous stipe) belonging to sect. Sicci ser. Leonini while M. cladophyllus (type variety) has very distinct caulocystidia, being referred amongst species of sect. Sicci ser. Atrorubentes. It is also uncertain whether the presence of broom cells in the stipitipellis may or may not be considered also one form of caulocystidium.

Additionally, setae like those found in the pileipellis of M. longisetosus, or like those in M. cohaerens, may also be found in lamellae and/or stipe in other species of sect. Sicci ser. Spinulosi. Clémençon (2012) explained that the term ‘setae’ has received a widened concept to accommodate setoid elements without the xanthocroic reaction, or having basal clamp connection, or with partly or completely hyaline wall, and to admit similar but richly branched cells, or similar, hair-like cells on the stipe and pileus of some agarics. In Marasmius, these elements were tentatively termed ‘spinulae’ by Clémençon (1982), but the term ‘setae’ was conserved (Antonín & Noordeloos 1993), only being categorized according to topography. Considering the setae in sect. Sicci ser. Spinulosi, it is difficult to disregard the fact that the long, setoid elements (at least, the majority) in M. atrorubens (also var. dumontii with elongate, hair-like structures in the pileipellis), called as caulocystidia (Fig. 7), can be considered setae.

The third case is the presence of pleurocystidia. There is certain difficulty to define true pleurocystidia among a) those sometimes called ‘gloeocystidia’ which are long and refractive cells, often with constrictions (i.e., M. ferrugineus, M. haematocephalus, M. hypophaeus, M. siccus); b) the non-refractive, reduced, but conspicuous elements (i.e., M. azectus, M. helvolus, M. hinnuleus, M. luteoolivaceus, M. spegazzinii, M. suthepensis); c) the inconspicuous elements or cystidioles in some species in ser. Leonini studied by Singer (1976) (such as M. bambusinus, M. bellus, M. berteroi, M. corrugatus, M. cuatrecasasii, M. helvoloides, M. pusio and M. neglectus and in M. hobbitii (in this study); and d) broom cells in the face of the lamellae (i.e., M. setulosifolius and M. pleurocanthus). According to D.J. Lodge (pers. comm.), the case of letter ‘c’ can be solved by testing the structures with Cresyl Blue and, if they are metachromatic unlike the basidia and basidioles, should be considered pleurocystidia. In this study, it was observed that the wide pleurocystidia concept (Singer 1958, 1976) is not informative to group species with these structures (whatever the form) in a single group based on phylogenetic analyses using ITS and LSU. Cases such as M. ambicellularis and M. linderioides and also the close relationship between M. hobbitii and M. luteoolivaceus (all discussed in the taxonomic section) also point to phylogenetic inconsistency regarding the presence/absence of pleurocystidia.

The pileipellis composed exclusively of smooth cells was traditionally considered diagnostic to form a section (sect. Globulares sensu Singer) likewise those species having Siccus-type broom cells in the pileipellis for sect. Sicci. However, classification based on the type of pileipellis cells is not supported by phylogenetic analyses (Tan et al. 2009, Wannathes et al. 2009, this study). Pileipellis made up by only smooth cells, or only Siccus-type broom cells, or with both are only useful for separating species, or at most, to group closely related species forming a stirps. Smooth cells mixed with broom cells in the pileipellis may denote an intermediate pattern between the species of sect. Sicci and the species of sect. Globulares sensu Singer (1986). However, there is no phylogenetic evidence that supports this conclusion. The evidence suggests multiple emergence of the different types of cells in the pileipellis within the single large group (sect. Globulares). Sometimes both types of cells occurring in the pileipellis may be interpreted as occasional when a minority of smooth cells are inconspicuously found among abundant broom cells, also representing broom cells not fully formed. But in species such as M. congregatus, M. corrugatus (as described by Singer (1976)), M. nexus, M. pseudoniveus (all varieties), M. pseudoniveoaffinis and M. ambicellularis, both type of dermatocystidia appear to be equally distributed or smooth cells are even dominant over the broom cells, possibly indicating that this pattern may also be useful for delimitation of species and varieties.