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Nayaka, Sanjeeva, Debnath, Ambikesh (2023): Cyphellostereum indicum (Hygrophoraceae), a new species of basidiolichen from India. Phytotaxa 603 (3): 271-279, DOI: 10.11646/phytotaxa.603.3.6, URL: http://dx.doi.org/10.1094/PDIS-04-22-0755-PDN

Neohygrocybe pseudoingrata, a new waxcap species known from Slovakia and the Czech Republic, is characterised by its pale greyish coloured and often robust basidiomata (or sporocarps), nitrous smell, context without colour changes, hollow, contorted and compressed stipe and smooth or slightly fibrillose pileus surface. Based on morphology and DNA analysis of ITS and LSU sequences of the collected specimens, N. pseudoingrata belongs to Neohygrocybe sect. Neohygrocybe together with N. ovina, N. nitrata and N. ingrata. Collections of N. pseudoingrata form a well-supported clade in phylogenetic trees.

Key to the known species of Hygrophorus subgenus Colorai from China 1. Cheilocystidia and pleurocystidia absent...........................................................................................................................................2 1. Cheilocystidia and pleurocystidia present..........................................................................................................................................3 2. Pileus surface dry, raw umber to pale ochre...................................................................................................................... H. robustus 2. Pileus surface glutinous, dark yellow or olivaceous brown...................................................................................... H. viridiflavidus 3. Floccules at the stipe apex absent.......................................................................................................................................................4 3. Floccules at the stipe apex present...................................................................................................................................................13 4. Stipe apex has an annular zone...........................................................................................................................................................5 4. Stipe apex has none annular zone.......................................................................................................................................................7 5. Pileus blackish-brown to brown...................................................................................................................................... H. annulatus 5. Pileus light yellow to yellow..............................................................................................................................................................6 6. Pileus plane, with a broad small umbo........................................................................................................................... H. esculentus 6. Pileus plane, with a depressed center................................................................................................................................ H. xiangjun 7. Pileus has reddish to purplish tinge....................................................................................................................................................8 7. Pileus has brown tinge......................................................................................................................................................................11 8. Pileus> 15 cm diam.; basidiospores small, 5.5–7 × 3.5–4.8 μm.................................................................................... H. orientalis 8. Pileus 9. Pileus> 10 cm diam.; associated with Fagaceae..............................................................................................................................10 9. Pileus 10. Pileus applanate, context thick (up to 2.4 cm.................................................................................................................. H. deliciosus 10. Pileus wavy and uplifted, context thin (0.4–0.9 cm.................................................................................................... H. qinggangjun 11. Pileus broadly convex or flat, slender basidia (41–57 × 6–10 μm)..................................................................................... H. alpinus 11. Pileus plane with a small umbo, shorter and wider basidia (33–45 × 8–11 μm................................................. H. pseudohypothejus 12. Basidiospores broad ellipsoid, Qm = 1.48, surface bacillate ornamentation.............................................................. H. parvirussula 12. Basidiospores narrow ellipsoid, Qm = 1.86, surface smooth...................................................................................... H. yunnanensis 13. Odor sweetish, reminiscent of bitter almonds..................................................................................................................................14 13. Odor indistinct..................................................................................................................................................................................15 14. Associated with Picea spp...................................................................................................................................... H. agathosmoides 14. Associated with Pinus spp............................................................................................................................................. H. pinophilus 15. Habitat in broad-leaved forests....................................................................................................................................... H. atrofuscus 15. Habitat in mixed broadleaf-conifer tree forests................................................................................................................................16 16. Pileus 4–7cm diam., yellowish-brown to brown, margin wavy and ruffled like the hem of a skirt........................... H. brunneiceps 16. Pileus 3–4.5 cm diam., applanate, margin usually uplifted at maturity; fawn-colored to yellowish-brown near the margin, brown at centre............................................................................................................................................................................... H. habaensis

Hygrophorus viridiflavidus, a new species from Shanxi Province in northern China, belonging to subgenus Colorati, section Limacini, is proposed in this study based on morphology and molecular systematics. Hygrophorus viridiflavidus is similar and closely related to H. glutinifer, but can be distinguished from the latter by its shorter and narrower basidia (35.0–46.5 × 8.0–10.5μm). Morphological characteristics and ITS-phylogeny support Hygrophorus viridiflavidus as a distinct species.

Hygrophorus viridiflavidus L. Fan & Y.X. Zhang sp. nov. (Fig. 2, 3) MycoBank:— MB846250 Diagnosis: differs from the other species of subgenus Colorati section Limacini in its medium-sized basidiomata, lamellae partially turning yellowish green when bruised, shorter and narrower basidia 35.0–46.5×8.0–10.5 μm, cystidioid elements absent, and longer and narrower basidiopores (8.5–)9.0–10.5(–11.0) × (4.5–)4.8–5.8(–6.0) μm. Etymology: viridiflavidus, viridi -, meaning green, flavidus, meaning yellowish, referring to the lamellae gradually turning light yellowish green when bruised. Holotype: CHINA. Shanxi Province, Qinshui County, Lishan Mountains, Zhongcun Forest Farm, on the ground in forest of Quercus spp., 112°1’27’’E, 35°29’7’’N, elev. 1,680 m, 24 August 2020, collected by H. Liu LH 1116 (BJTC FM978, GenBank Acc. No.: ITS = OP699731, nrLSU = OP699736, tef1-α = OP723351). Description: Basidiomata medium-sized, firm, fleshy, with dark greenish tone. Pileus 41–57(–98) mm, hemispherical or obtusely conical with strongly involute margin at first, then expanded to almost flat with prominent obtuse umbo, first rather dark yellow (#917625) or olivaceous brown (#676700) with dark greyish (#1b1b00) to blackish brown (#424200) centre, then paler especially in marginal zone towards light greyish (#786e4c) or olivaceous brown (#676700), mostly with olivaceous (#9b8957) or dark greenish (#4e4e00) tinge, centre always much darker, very strongly glutinous. Lamellae adnate to subdecurrent, thick and waxy, white (#ffffff), partly gradually turning light yellowish green (#cccc00) to drying cream (#ffeb07) when bruised. Stipe 38–79 × 10–23 mm, cylindro-clavate to ventricose, often flexuous or bent and tapering-rooting towards the base, connected with margin of cap by a glutinous hyaline veil when young, then veil burst and disappear with maturity, with an unconspicuous annular zone at the apex, pale yellowish white (#ffffd6), smooth or with white (#ffffff) floccules above the annular zone, covered by successive bands of thickly glutinous, dark brown (#856731) or greyish brown (#433419) flocci below, drying greybrown (#686042). Context thick, pure white (#ffffff), in stem also white or slightly yellowish at base. Smell indistinct. Taste not recorded. Spore deposit white. Basidiopores ellipsoid to broadly ellipsoid or somewhat ovoid, measuring (8.5–)9.0–10.5(–11.0) × (4.5–)4.8–5.8(– 6.0) μm; Q =1.58–2; Qm =1.79±0.11, with an oblique hilar appendage 1–2 μm long; inamyloid, hyaline and smooth or microguttulate in water, appearing monoguttulate in KOH. Basidia 2-spored or 4-spored, clavate, 35.0–46.5 × 8.0–10.5 μm, some with oily refractive content in KOH; sterigmata 3.5–5.5 μm long. Cystidioid elements absent. Hymenophoral trama divergent, comprised of cylindrical, interwoven, branched hyphae 3–13 μm wide. Pileipellis an ixotrichoderm composed of thick-walled, hyaline or brown-pigmented interwoven hyphae 2.5–4 μm wide, hyaline or with olive green intracellular pigment, sparsely branching and embedded in a gelatinous matrix; incrustations or extracellular pigment not seen. Subpellis composed of more or less parallel hyphae up to 9 μm wide. Stipitipellis an ixotrichoderm, made up of branched hyphae 3–5 μm wide, with yellowish brown intracellular pigment; terminal elements rounded to attenuated, not enlarged. Stipititrama composed of densely packed, more or less parallelly arranged, subhyaline hyphae up to 9 μm wide. Floccules at the apex composed of compact fascicles of branched hyphae with cylindrical or slender clavate terminal elements 37–74 × 5–9 μm. Clamp connections present in all tissues. Ecology & Distribution: associated with Quercus spp. in elevation 1,680 m, only known from Shanxi Province, northern China. Additional specimens examined (paratypes): CHINA. Shanxi Province, Qinshui County, Lishan Mountains, Zhongcun Forest Farm, on the ground in forest of Quercus spp., 112°1’21’’E, 35°29’3’’N, elev. 1,680 m, 24 August 2020, collected by H. Liu LH 1139 (BJTC FM999, GenBank Acc. No.: ITS = OP699730, nrLSU = OP738418, tef1-α = OP723352)., Published as part of Zhang, Yu-Xin, Mao, Ning & Fan, Li, 2023, Hygrophorus viridiflavidus sp. nov. (Hygrophoraceae, Agaricales), from North China, pp. 187-197 in Phytotaxa 579 (3) on pages 191-192, DOI: 10.11646/phytotaxa.579.3.4, http://zenodo.org/record/7550376

Hygrocybe calciphila, an uncommon species of the genus was found for the first time in Southeastern Europe in forests of Pinus heldreichii in Bulgaria. Description and illustrations of the finding are presented. The European distribution of the species is summarized and briefly discussed, with emphasis on the scarce previous findings in Southern Europe.

8) Dictyonema umbricola V. Marcano, sp. nov.; (Fig. 11) Mycobank MB#838498 Diagnosis:— Sicut Dictyonema discocarpum, sed filamentis furcatis, marginibus albidis, et hyphis fibulatis differt. Thallus muscicola. Hyphothallus praesentia. Hymenocarpus elevatus, disciformibus, margine tomentoso. Hymenium N non reagens. Basidiosporae obovatae, 9–10 μm longae, 4–5 μm crassae. Type:— VENEZUELA. Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua cloud forests, elev. 2550 m; 15 June 2020, V. Marcano & L. Castillo 20-91 (holotype MER, isotype B, private herb. V. Marcano). Etymology:—The epithet refers to the very shady habitat. Description:— Thallus growing on bryophytes, filamentous, dark blue-green, covering shorter areas of the substrate, by irregular, discontinuous patches up to 12 cm across, composed of loosely interwoven, dark green to bluish green fibrils leaving interspaces and bordered by a distinct, discontinuous, white margin (prothallus). Fibrils subascending or horizontal, with whitish apices, broad, more loosely and irregularly arranged and leaving interspaces. Thallus (when fresh) in section 1.25–2.5 mm thick, composed of a thick, loose, photobiont (Rhizonema) layer (0.75–1.75 mm) and a pale brown medulla (0.5–0.75 mm) (hypothallus); photobiont layer composed of cyanobacterial filaments covered by thin elongated fungal cells, forming jigsaw-puzzle-shaped cells; cyanobacterial filaments branched, composed of 13–17 μm wide and 7–10 μm high, bluish green cells, cylindrical, in chains, penetrated by tubular fungal hyphae (haustoria); heterocytes frequent, pale to slightly bright yellow, 12–13 μm wide and 2–3 μm high, narrowly elliptic; cells of hyphal sheath variably wavy in lateral outline, 2–3 μm thick; hyphae of hypothallus straight, branched, hyaline, smooth, 3–5 μm thick. Clamp connections present on the hyphae, abundant. Hymenophore (when fresh) developed as small, rounded to irregular, corticioid patches, dispersed on the upper surface of thallus and resembling apothecial discs, 0.6–2 mm diam., with pale yellowish, smooth surface and white, minutely tomentose margins, 125–200 μm diam.; hymenophore in section 50–120 μm thick, composed of a paraplectenchymatous layer connected to loose medullary hyphae, 5–6 μm wide; hymenium composed of numerous, palisade-like basidioles and scattered basidia; basidiospores (only a few seen) obovate, non-septate, hyaline, 9–10 × 4–5 μm. Chemistry:—Thallus K–, C–, KC–, P–; medulla K–, C–, KC–, P–, N –, ER–; basidiocarp K–, C–, KC–, P–, N –, ER– (n = 8). No lichen compounds detected by TLC. Distribution and Ecology:—This species is known only from the well-developed type collection. It was found growing on mosses in a very shady and humid rainforest at elev. 2000–2650 m (Fig. 9A). Remarks:— Dictyonema umbricola is characterized by horizontal or subascending fibrils, branched cyanobacterial filaments, jigsaw-puzzle-shaped cells, abundant clamp connections on the hyphae and a negative N spot test.This species resembles D. discocarpum Lücking, Dal Forno & Wilk (Lücking et al. 2013b: 7). Both species have a hymenophore composed of disc-shaped patches resembling apothecial discs, with pale yellowish, smooth surface, minutely tomentose margins and jigsaw-puzzle-shaped cells. However, the latter has shelf-like filamentous thallus, a hymenophore in smaller disc-shaped patches (0.3–0.5 mm diam.) dispersed on the underside of the thallus, a hypothallus with byssoid margin, and lacks clamp connections. Dictyonema discocarpum is known only from Bolivia (Lücking et al. 2013b). Some species of the genus Cyphellostereum (Lawrey et al. 2009) have disc-like hymenophores forming resupinate, rounded patches, viz., C. jamesianum Dal Forno & Kaminsky (Dal Forno et al. 2019: 338), and C. georgianum Dal Forno, McMullin & Lücking (Dal Forno et al. 2019: 337). Yet, these species are characterized by a crustose, dark green thallus formed by loose individual fibrils, a fungal sheath consisting of irregular, thin, branched hyphae surrounding the cells, and absence of jigsaw-puzzle-shaped cells. Both species are known only from the southeastern United States and Puerto Rico. Additional specimens examined (paratypes):— Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua, cloud forests, elev. 2650 m; 12 October 2020, V. Marcano & L. Castillo 20-577 (MER, private herb. V. Marcano); Parque Sierra Nevada de Mérida, road towards Lake Coromoto, cloud forest, elev. 2000 m, V. Marcano & L. Castillo 21-110 (MER, private herb. V. Marcano)., Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on pages 218-220, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966, {"references":["Lucking, R., Dal Forno, M., Wilk, K. & Lawrey, J. D. (2013 b) Three new species of Dictyonema (lichenized Basidiomycota: Hygrophoraceae) from Bolivia. Acta Nova 6: 4 - 16.","Lawrey, J. D., Lucking, R., Sipman, H. J. M., Chaves, J. L., Redhead, S. A., Bungartz, F., Sikaroodi, M. & Gillevet, P. M. (2009) High concentration of basidiolichens in a single family of agaricoid mushrooms (Basidiomycota: Agaricales: Hygrophoraceae). Mycological Research 113: 1154 - 1171. https: // doi. org / 10.1016 / j. mycres. 2009.07.016","Dal Forno, M., Kaminsky, L., Rosentreter, R., McMullin, R. Troy, Aptroot, A. & Lucking, R. (2019) A first phylogenetic assessment of Dictyonema s. lat. in southeastern North America reveals three new basidiolichens, described in honor of James D. Lawrey. Plant and Fungal Systematics 64 (2): 383 - 392. https: // doi. org / 10.2478 / pfs- 2019 - 0025"]}

4) Dictyonema duidense V. Marcano , sp. nov.; (Fig. 7) Mycobank MB#838342 Diagnosis:— Sicut Dictyonema barbatum sed hymenocarpis resupinatis, irregularis differt. Thallus corticola, byssaceus, crassus, pagina areolata, rugosa, hyphis non fibulatis. Hymenium N non reagens. Sporae guttaeformes, hyalinae, non septatae, 7–8 μm longae, 2–4 μm latae. Type:— VENEZUELA. Amazonas: Alto Orinoco, summit northern Cerro Duida, growing on tree trunks at margin of small tepuian forests, elev. 2100 m, March 1996, L. Galiz & V. Marcano AMA-757 (holotype MER, isotype B, private herb. V. Marcano). Etymology:—The epithet refers to the locality where the type specimens were collected. Description:— Thallus shelf-like, corticolous, byssaceous, filamentous, epiphytic on tree trunks, covering larger areas of the substrate with a thick crust, in irregular, confluent patches up to 35 cm across, composed of densely arranged, dark aeruginous fibrils with distinct interspaces resting on a thick, byssoid, irregularly interwoven medulla (hypothallus), visible as narrow, 0.8–1.1 mm wide marginal zone, strongly contrasting with the aeruginous fibrils. Lobes projecting vertically from the substrate, semicircular, coated, dark blue-green, very broad, 15–25 cm; surface areolate, rugose, with entire, white margins, 1–1.5 cm wide, lacking photobiont filaments; photobiont layer composed of numerous cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsaw-puzzle-shaped cells; cyanobacterial filaments not branched, composed of 20–25 μm wide and 4–6 μm high, bluish green cells penetrated by tubular fungal hyphae (haustoria). Thallus in section 3–6(–7) mm thick, not forming a distinct photobiont layer and medulla but instead composed of aeruginous fibrils connected at base (underside) to a pale brown, loosely woven hypothallus. Clamp connections absent. Hymenophore forming irregular, reticulated, resupinate patches dispersed on the underside, with entire margin, whitish, smooth surface; hymenophore composed of a paraplectenchymatous layer connected to loose medullary hyphae; hymenium composed of numerous, palisade-like basidioles and scattered basidia; basidiospores ellipsoid to narrowly drop-shaped, non-septate, hyaline, 7–8 × 2–4 μm. Chemistry:—Thallus K–, C–, KC–, P–; medulla K–, C–, KC–, P–, N –, ER–; basidiocarp K–, C–, KC–, P–, N –, ER– (n = 6). No lichen compounds detected by TLC. Distribution and Ecology:— Dictyonema duidense is found growing as an epiphyte on tree bark in altotepuian forests at 1500–2100 m. It is known only from the type locality. Remarks:— Dictyonema duidense is another species in the complex formerly recognized as the single species, Dictyonema sericeum (Parmasto 1978). In their treatment of Dictyonema s. str., Lücking et al. (2013a,b) indicated that the phenotype usually identified with the name D. sericeum is a complex of several, phylogenetically distinct species, differing morphologically in the orientation and arrangement of the fibrils and in the morphology of the hymenophore. According to Lücking et al. (2014b) the thallus of D. sericeum type is much more compacted than any of the species collected in South America, indicating, along with the geographic range, that D. sericeum s.str. is a distinct taxon possibly restricted to the Caribean. Dictyonema duidense is characterized by a shelf-like filamentous thallus, with thick, very broad lobes, areolate, rugose surface and lacking clamp connections. It is very similar to D. barbatum Dal Forno, Bungartz & Lücking (Dal Forno et al. 2017: 62) from the Galápagos Islands (Dal Forno 2015, Dal Forno et al. 2017). Both species have a shelflike filamentous thallus and lack clamp connections on the hyphae but D. barbatum is distinguished by the color of the prothallus (old specimens with a darkened hue, yellow to light brown) and the continuous hymenophore. Dictyonema duidense resembles D. excentricum C. Agardh (Kunth 1822: 1). This latter is characterized by thick, horizontally arranged bundles of fibrils and is known only from French Guiana. Additional specimens examined (paratypes):— Amazonas: summit northern Cerro Duida, growing on tree bark in small tepuian forests, elev. 1500 m, March 1996, L. Galiz & V. Marcano AMA-539, 540 (TFAZ, VEN, private herb. V. Marcano)., Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on pages 212-213, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966, {"references":["Parmasto, E. (1978) The genus Dictyonema (' Thelephorolichenes'). Nova Hedwigia 29: 99 - 144.","Lucking, R., Dal Forno, M., Lawrey, J. D., Bungartz, F., Rojas, M. E. H., Hernandez M., J. E., Marcelli, M. P., Moncada, B., Morales, E. A., Nelsen, M. P., Paz, E., Salcedo, L., Spielmann, A. A., WilK, K., Will-Wolf, S. & Yanez-Ayabaca, A. (2013 a) Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales: Hygrophoraceae), with a key to all accepted genera and species in the Dictyonema clade. Phytotaxa 139: 1 - 38. https: // doi. org / 10.11646 / phytotaxa. 139.1.1","Lucking, R., Dal Forno, M., Sikaroodi, M., Gillevet, P. M., Bungartz, F., Moncada, B., Yanez-Ayabaca, A., Chaves, J. L., Coca, L. F. & Lawrey, J. D. (2014 b) A single macrolichen constitutes hundreds of unrecognized species. Proceedings of the National Academy of Sciences of the United States 111: 11091 - 11096. https: // doi. org / 10.1073 / pnas. 1403517111","Dal Forno, M., Bungartz, F., Yanez-Ayabaca, A., Lucking, R. & Lawrey, J. D. (2017) High levels of endemism among Galapagos basidiolichens. Fungal Diversity 85: 45 - 73. https: // doi. org / 10.1007 / s 13225 - 017 - 0380 - 6","Dal Forno, M. (2015) Evolution and Diversity of the basidiolichen clade Dictyonema (Agaricales: Hygrophoraceae). PhD Dissertation, pp. 1 - 294.","Kunth, C. S. (1822) Synopsis plantarum, quas in itinere circa plagas Orbis Novi colleg. Humboldt et Bonpland. Paris, pp. 1 - 491."]}

7) Dictyonema subsericeum V. Marcano , sp. nov.; (Fig. 10) MycoBank MB#838497 Diagnosis:— Sicut Dictyonema sericeum, sed sed hyphis fibulatis differt. Thallus corticola, byssaceus, crassus, pagina hirsuta. Lobis spongiosis, maxime variantibus statura magnitudine latitudine colore et forma. Hymenocarpus resupinatus, hapteriformi. Hymenium N non reagens. Sporis guttaeformibus, hyalinis, non septatis, 4.5–5.5 μm longis, 3.7–4.7μm latis. Type:— VENEZUELA. Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua cloud forests, elev. 2650 m; 5 June 2020, V. Marcano & L. Castillo 20-84 (holotype MER, isotype B, private herb. V. Marcano). Etymology:—The epithet refers to the thallus appearing closely related to the species D. sericeum. Description:— Thallus corticolous, byssaceous, filamentous, forming a thick crust, epiphytic on tree branches. Thallus surface plane or irregular, projecting horizontally from the substrate, covering up to 20 cm diameter, composed of numerous imbricate to partially fused lobes. Lobes projecting parallel from the substrate, semicircular or hemispherical, variable in color, size and shape, when small forming white, filamentous finger-like outgrowths, 4–7 mm broad, when large semicircular, 18–21 mm broad, greenish blue in the bottom side with fibrillose, white margins, 4–5.5 mm broad, and when located in the center of the thallus, brown, forming hemispherical clusters, 5–7(–8) mm broad; surface hirsute. Thallus (when fresh) in section 3–5.8 mm thick, composed of an appressed, pale brown medulla (1.6–2.5 mm) (hypothallus), a broad, very loose, aeruginous photobiont (Rhizonema) layer (1–3 mm) and frequently with an upper layer (0.5–1.25 mm) composed of irregularly arranged, long, white fibrils, lacking photobiont filaments; photobiont layer composed of numerous cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsawpuzzle-shaped cells; cyanobacterial filaments not divided, composed of 22–27 μm wide and 4.5–7(–8) μm high, bluish green cells penetrated by tubular fungal hyphae (haustoria); heterocytes frequent, bright yellow, elliptic, 13–18 μm wide and 5–8 μm high; hyphae associated with hyphal sheath straight, hyaline, 5–6 μm thick, with clamp connections, finely papillose just before the apices. Hymenophore (when fresh) developed as small, angular to elongate, white, resupinate patches dispersed on the underside and resembling irregular, large (500–1200 μm wide) attachment hapters with white, more or less smooth or pruinose surface, and fibrillose margins; hymenophore composed of a paraplectenchymatous layer connected to loose medullary hyphae, 6–7.5 μm wide; hymenium composed of numerous, palisade-like basidioles and scattered basidia; basidia 28–37 × 4–6 μm, 4-sterigmate; basidiospores globose or subglobose, non-septate, hyaline, 4.5–5.5 × 3.7–4.7 μm. Chemistry:—Thallus K–, C–, KC–, P–; medulla K–, C–, KC–, P–, N –, ER–; basidiocarp K–, C–, KC–, P–, N –, ER– (n = 8). No lichen compounds detected by TLC. Distribution and Ecology:— Dictyonema subsericeum was found growing as an epiphyte on branches in the canopy of rainforests at elev. 2200–2250 m (Fig. 8A). It is known only from the well-developed type collection. Remarks:—Molecular phylogenetic data clearly show that D. sericeum sensu Parmasto contains many different species and even the shelf-like forms representing D. sericeum in a narrow sense are more than one species (Dal Forno et al. 2013). Dictyonema subsericeum is characterized by a thallus forming a thick crust, filamentous lobes, variability of color (white, greenish blue, brown), size and shape (semicircular and hemispherical), with a hirsute surface, fibrillose margins, clamp connections on the hyphae, a hymenophore resembling large, irregular attachment hapters with a white surface and a negative N spot test. Dictyonema subsericeum is like D. sericeum (Dal Forno et al. 2013), D. hapteriferum and D. spongiosum Berkeley & M. A. Curtis (1868: 335), the latter representing the lineage labeled D. sericeum 3 in Dal Forno et al. (2013). These species are characterized by semicircular, shelf-like, filamentous lobes and absence of clamp connections. Dictyonema hapteriferum has aeruginous fibrils bordered by a narrow, byssoid, white margin, a hymenophore resembling attachment hapters with beige, smooth surface. Dictyonema spongiosum has a thick, spongiose upper surface composed of bundles of vertically projecting fibrils and it is known only from Cuba and Guatemala. Dictyonema subsericeum might also be confused with D. irpicinum and D. ligulatum (Krempelhuber 1875: 15) Zahlbruckner (1908: 239). Both species have a lobed surface mottled white and blue-green with coarse, finger-like outgrowths, and hyphae with clamp connections but they are restricted to the Paleotropics (Lücking et al. 2013a, Dal Forno 2015, Dal Forno et al. 2019). Dictyonema subsericeum is like to D. album Lücking & Timdal (2016: 192) from Mauritius. However, this latter species has a combination of irregularly arranged tufts of fibrils that do not form a distinctly layered thallus, being dominated by straight hyphae with very few associated cyanobacterial fibrils, giving the thallus a whitish appearance and it lacks clamp connections on the hyphae (Lücking & Timdal 2016). Additional specimens examined (paratypes):— Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua, cloud forests, elev. 2650 m; V. Marcano & L. Castillo no. 20-85 (MER, private herb. V. Marcano)., Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on pages 216-218, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966, {"references":["Dal Forno, M., Lawrey, J. D., Sikaroodi, M., Bhattarai, S., Gillevet, P. M., Sulzbacher, M. & Lucking, R. (2013) Starting from scratch: evolution of the lichen thallus in the basidiolichen Dictyonema (Agaricales: Hygrophoraceae). Fungal Biology 117: 584 - 598. https: // doi. org / 10.1016 / j. funbio. 2013.05.006","Krempelhuber, A. von (1875). Lichenes quos legit O. Beccari in insulis Borneo et Singapore annis 1866 et 1867. Nuovo Giornale Botanico Italiano 7: 5 - 67.","Zahlbruckner, A. (1908) Flechten (Lichenes). In: Engler, H. G. & Prantl, K. A. (Eds.) Die Naturlichen Pflanzenfamilien I (1): 1 - 239.","Lucking, R., Dal Forno, M., Lawrey, J. D., Bungartz, F., Rojas, M. E. H., Hernandez M., J. E., Marcelli, M. P., Moncada, B., Morales, E. A., Nelsen, M. P., Paz, E., Salcedo, L., Spielmann, A. A., WilK, K., Will-Wolf, S. & Yanez-Ayabaca, A. (2013 a) Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales: Hygrophoraceae), with a key to all accepted genera and species in the Dictyonema clade. Phytotaxa 139: 1 - 38. https: // doi. org / 10.11646 / phytotaxa. 139.1.1","Dal Forno, M. (2015) Evolution and Diversity of the basidiolichen clade Dictyonema (Agaricales: Hygrophoraceae). PhD Dissertation, pp. 1 - 294.","Dal Forno, M., Kaminsky, L., Rosentreter, R., McMullin, R. Troy, Aptroot, A. & Lucking, R. (2019) A first phylogenetic assessment of Dictyonema s. lat. in southeastern North America reveals three new basidiolichens, described in honor of James D. Lawrey. Plant and Fungal Systematics 64 (2): 383 - 392. https: // doi. org / 10.2478 / pfs- 2019 - 0025","Lucking, R. & Timdal, E. (2016) New species of Dictyonema and Cyphellostereum (lichenized Basidiomycota: Hygrophoraceae) from tropical Africa and the Indian Ocean, dedicated to the late Hildur Krog. Willdenowia 46: 191 - 199. https: // doi. org / 10.3372 / wi. 46.46115"]}

A study of Acantholichen P.M. Jørgensen, Cyphellostereum D.A. Reid and Dictyonema s.str. C. Agardh ex Kunth from the Venezuelan Andes (Sierra Nevada de Mérida) and the Guayana Highlands (Alto Orinoco) revealed the presence of eight species new to science, viz., Acantholichen dendroideus V. Marcano, Cyphellostereum mucuyense V. Marcano, Dictyonema andinum V. Marcano, D. duidense V. Marcano, D. laurae V. Marcano, D. subinvolutum V. Marcano, D. subsericeum V. Marcano and D. umbricola V. Marcano. Acantholichen dendroideus grows on mosses in shady and humid disturbed forest and is characterized by densely and dichotomously branched microsquamules, with a pruinose surface and large acanthohyphidia. Cyphellostereum mucuyense grows on mosses and sandy soils, lacks a jigsaw-puzzle-shaped hyphal sheath and prothallus, the fibrils are not penetrated by tubular fungal hyphae and the basidiocarp surface is finely arachnoid. This is the first record of Cyphellostereum in Venezuela. Dictyonema andinum might be confused with D. irrigatum (Berkeley & M. A. Curtis) Lücking from the Andaman Islands, but differs in the broader and ascending fibrils, a thicker photobiont layer and hypothallus, as well as the formation of stereoid-corticioid hymenophores. Dictyonema duidense from the Guayana Highlands and D. subsericeum from the Venezuelan Andes belong in the group of shelf-like morphotypes, with a semicircular thallus resembling D. sericeum s.str. Dictyonema duidense grows on tree trunks and is characterized by thick, very broad lobes and an areolate, rugose surface and it lacks clamp connections whereas D. subsericeum grows on tree branches and is characterized by variability of color (white, greenish blue, brown), size and shape (semicircular to hemispherical), filamentous lobes, a hirsute surface, fibrillose margins, clamp connections on the hyphae, cyphelloid hymenophores resembling attachment hapters, and very small, globose to subglobose basidiospores. Dictyonema laurae resembles D. subsericeum but is characterized by a continuous greenish blue crust-like thallus with an irregular surface, with discontinuous, fibrillose, and narrow margins projecting and ascending from the substrate, a hapteriform hymenophore, and its lacks clamp connections on the hyphae. Dictyonema subinvolutum is characterized by a jigsaw-puzzle-shaped hyphal sheath, fibrils penetrated by tubular fungal hyphae, and a smooth, pruinose, resupinate hymenophore. Finally, D. umbricola grows on mosses in very shady habitats and produces a corticioid hymenophore raised from the thallus, resembling apothecial discs, jigsaw-puzzle-shaped cells around the filaments and a distinct, discontinuous, white prothallus. Of the eight species of Dictyonema s.lat. registered from Venezuela, seven are known only from the Venezuelan Andes (A. dendroideus, C. mucuyense, D. andinum, D. laurae, D. subsericeum, D. subinvolutum, D. umbricola) and one is known only from the Venezuelan Amazonas (D. duidense). The high concentration of Acantholichen, Cyphellostereum and Dictyonema species in tropical America and especially of the latter genus in the Venezuelan Andes indicates that this region constitutes an important centre for speciation and endemism of Dictyonema clade/subtribe Dictyonematinae.

1) Acantholichen dendroideus V. Marcano , sp. nov.; (Fig. 4) Mycobank MB#838340 Diagnosis:— Sicut Acantholichen campestris, sed squamis dense dichotome furcatis differt. Thallus muscicola, microsquamulosus. Squamulae microfruticosae, pruinosae. Soredia nulla. Hyphis tegentibus fibrillas quadratorhombicis, non fibulatis. Acanthohyphidiae 20–27 longae × 5–7 μm crassae, subglobosae vel pyriformae, conspicue spinosae, abundantes. Etymology:—The epithet refers to the characteristic squamules which are divided at the top fine ramifications. Type:— VENEZUELA. Mérida: Parque Sierra Nevada de Mérida, La Mucuy Alta, elev. 2150 m; 5 November 2020, V. Marcano 20-901 (holotype MER, isotype B, private herb. V. Marcano). Description:— Thallus microsquamulose; squamules thin and mostly elongated, 1–2 mm long, 100–180 μm broad, microfruticulose, many growing together, occasionally thus shading one another, olivaceous when fresh, light blue grey when dry, the shaded parts becoming necrotic and pale beige, attached basally to the substrate, densely and dichotomously divided from the tips; microbranches swollen, 1–3(–4) times divided, coral-like, ascendent or erect, 300–350 μm long, 40–60 μm broad. An overall uniformly white-pruinose appearance due to acanthohyphidia. Soredia absent. Thallus in cross section 60–110 μm thick, composed of a thick photobiont (Rhizonema) layer (60–80 μm), a very thin paraplectenchymatous cortex and pale brown medulla. Photobiont forming clusters of coiled cyanobacterial filaments (chroococcoid; 7–14 μm diam.) inside the thallus microsquamulose, wrapped in a dense hyphal sheath formed by jigsaw-puzzle-shaped cells. Acanthohyphidia variable in size, usually small (4–8 μm × 4–5 μm), irregular, subglobose to pyriform, sometimes large [20–24(–27) × 5–7 μm] and then elongate, apically thickened, conspicuously spiny, abundant, present on upper and lower surfaces. Clamp connections not observed. Chemistry:—Thallus K–, C–, KC–, P–; medulla K–, C–, KC–, P–, N + (pink) (n = 6; V. Marcano). No lichen compounds detected by TLC. Distribution and Ecology:—This species is known only from the well-developed type collection. It was found on mosses in a humid, shady disturbed forest growing on Persea americana P. Miller (1768: HW’95) associated with Cora, Pseudoparmelia Lynge (1914: 15) and Usnea species at 2100–2550 m (Fig. 3A). Remarks:—The genus Acantholichen, previously thought to be monospecific (Lawrey et al. 2009), currently includes six species, viz., A. sorediatus Dal Forno, Sipman & Lücking (2016: 49) and A. pannarioides from Costa Rica (this latter also from Venezuela) (Jørgensen 1998), A. albomarginatus Dal Forno, Marcelli & Lücking (2016: 43) and A. campestris Dal Forno, Spielmann & Lücking (2016: 44) from Brazil, A. galapagoensis Dal Forno, Bungartz & Lücking (2016: 44) from the Galápagos islands, and A. variabilis Dal Forno, Coca & Lücking (2016: 51) from Colombia (Dal Forno 2015, Dal Forno et al. 2016, 2017). Acantholichen dendroideus is morphologically distinct from all other known Acantholichen species. It is characterized by a light blue-grey (when dry) thallus, ascendent, dichotomously and densely branched (from the tips) microsquamules, a pruinose surface and large acanthohyphidia. The new species somewhat resembles A. campestris from Brazil and A. galapagoensis from the Galápagos islands. Both species lack soredia and have a branched thallus and pyriform acanthohyphidia. However, in A. campestris the squamules are not dichotomously branched, are grey (when dry), and show a microfruticulose appearance while growing into vertical structures; the acanthohyphidia are small (10–15 × 6–8 μm). It is the only species known that grows on rocks. In contrast A. galapagoensis has dark olivaceous grey (when dry), elongated, intricate and tiled squamules, a non-fruticose appearance, small to medium acanthohyphidia (12–16 × 6–10 μm), and grows on mosses and liverworts (Dal Forno et al. 2016). Additional specimens examined (paratype):— Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua, cloud forests, elev. 2550 m; June 2020, V. Marcano & L. Castillo no. 20-416 (MER, private herb. V. Marcano)., Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on pages 207-208, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966, {"references":["Miller, P. (1768) The Gardeners Dictionary. Eight edition, s. n .. John & Francis Rivington, London, 3 volumes (without pagination).","Lynge, B. A. (1914) Die Flechten der ersten Regnelischen Expedition. Die Gattungen Pseudoparmelia gen. nov. und Parmelia Ach. Arkiv fOr Botanisk 13 (13): 1 - 172.","Lawrey, J. D., Lucking, R., Sipman, H. J. M., Chaves, J. L., Redhead, S. A., Bungartz, F., Sikaroodi, M. & Gillevet, P. M. (2009) High concentration of basidiolichens in a single family of agaricoid mushrooms (Basidiomycota: Agaricales: Hygrophoraceae). Mycological Research 113: 1154 - 1171. https: // doi. org / 10.1016 / j. mycres. 2009.07.016","Dal Forno, M., Lucking, R., Bungartz, F., Yanez-Ayabaca, A., Marcelli, M., Speilmann, A. A., Coca, L. F., Chaves, J. L., Aptroot, A., Sipman, H. J. M., Gillevet, P. & Lawrey, J. D. (2016) From one six: Unrecognized species diversity in the genus Acantholichen (lichenized Basidiomycota: Hygrophoraceae). Mycologia 108 (1): 38 - 55. https: // doi. org / 10.3852 / 15 - 060","Jorgensen, P. M. (1998) Acantholichen pannarioides, a new basidiolichen from South America. The Bryologist 101: 444 - 447. https: // doi. org / 10.2307 / 3244185","Lucking, R. & Timdal, E. (2016) New species of Dictyonema and Cyphellostereum (lichenized Basidiomycota: Hygrophoraceae) from tropical Africa and the Indian Ocean, dedicated to the late Hildur Krog. Willdenowia 46: 191 - 199. https: // doi. org / 10.3372 / wi. 46.46115","Dal Forno, M. (2015) Evolution and Diversity of the basidiolichen clade Dictyonema (Agaricales: Hygrophoraceae). PhD Dissertation, pp. 1 - 294.","Dal Forno, M., Bungartz, F., Yanez-Ayabaca, A., Lucking, R. & Lawrey, J. D. (2017) High levels of endemism among Galapagos basidiolichens. Fungal Diversity 85: 45 - 73. https: // doi. org / 10.1007 / s 13225 - 017 - 0380 - 6"]}

Key to Acantholichen, Cyphellostereum and Dictyonema s.str. in Venezuela 1a. Thallus microsquamulose to microfruticulose, pruinose; squamules densely and dichotomously branched from the tips; photobiont forming clusters of short, irregularly coiled threads (chroococcoid) inside the thallus microsquamulose; soredia absent; upper and lower surfaces in microscope view forming apically thickened, distinctly spiny, pyriform to subglobose, variable in size, sometimes elongate hyphae (acanthohyphidia); clamp connections absent............................................. Acantholichen dendroideus 1b. Thallus composed of distinct fibrils including cyanobacterial filaments (Rhizonema); acanthohyphidia absent; clamp connections present or absent................................................................................................................................................................................2 2a. Thallus directly on bark of trunks or branches; lobes semicircular and/or hemispherical, variable in color, size and shape; basidiocarp N–; Venezuelan Andes or Guayana Highlands................................................................................................................................ (3) 2b. Thallus on mosses, liverworts or soil, appressed filamentous or loose and not forming semicircular lobes; basidiocarp N+ or N–; Venezuelan Andes (Sierra Nevada de Mérida)............................................................................................................................... (4) 3a (2). Thallus on tree branches; lobes projecting parallel from the substrate, semicircular and hemispherical, variable in color, size and shape; surface hirsute; clamp connections present; basidiocarp with resupinate patches dispersed on the underside and resembling large, irregular attachment, hapters, margin fibrillose; Venezuelan Andes (Sierra Nevada de Mérida)..... Dictyonema subsericeum 3b. Thallus on tree trunk, shelf–like filamentous, forming semicircular lobes, projecting upright from the substrate, surface areolate, rugose; clamp connections absent; basidiocarps forming irregular, reticulate, resupinate patches dispersed on the underside, margin entire; Guayana Highlands (Cerro Duida, Alto Orinoco).................................................................................. Dictyonema duidense 4a (2). Hymenophore resupinate; jigsaw-puzzle-shaped hyphal sheath present or absent; prothallus absent; clamp connections present or absent.............................................................................................................................................................................................. (5) 4b. Hymenophore corticioid; jigsaw-puzzle-shaped hyphal sheath present; prothallus present; clamp connections present.............. (6) 5a (4). Jigsaw-puzzle-shaped hyphal sheath absent; fibrils not penetrated by tubular fungal hyphae haustoria, horizontally arranged and closely appressed; cyanobacterial filaments 8–10 μm wide; clamp connections present; basidiocarp surface finely arachnoid, N–................................................................................................................................................................ Cyphellostereum mucuyense 5b. Jigsaw-puzzle-shaped hyphal sheath present; fibrils penetrated by tubular fungal hyphae haustoria, ascendent and loosely arranged; cyanobacterial filaments> 10 μm wide; clamp connections present or absent; basidiocarp surface smooth, N+ (pink)......................................................................................................................................................................................................... (7) 6a (4). Fibrils irregularly arranged, ascending or erect; cyanobacterial filaments not divided, 20–25 μm wide; hymenophores forming stereoid-corticioid patches, upper surface smooth, lower surface pruinose, spongy, N+ (red)......................... Dictyonema andinum 6b. Fibrils horizontal or subascending; cyanobacterial filaments divided, 13–17 μm wide hymenophores raised from the thallus, resembling apothecial discs, surface pale yellowish, smooth, margins white, minutely tomentose; N.......... Dictyonema umbricola 7a (5). Thallus developing a mat of interwoven fungal–cyanobacterial fibrils, leaving interspaces; cyanobacterial filaments 12–15 μm wide; clamp connections present; basidiocarp developed as effuse patches, with pruinose surface, margins slightly involute............................................................................................................................................................................. Dictyonema subinvolutum 7b. Thallus forming a thin crust (1.75–2 mm), bordered by a discontinuous, narrow, white margin projecting vertical from the substrate; cyanobacterial filaments 12–17 μm wide; clamp connections absent; basidiocarp hapteriform, with finely reticulate surface........................................................................................................................................................................................ Dictyonema laurae, Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on page 220, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966

5) Dictyonema laurae V. Marcano , sp. nov.; (Fig. 8) Mycobank MB#840644 Diagnosis:— Sicut D. subsericeum, sed hyphis non fibulatis differt. Thallus muscicola, byssaceus, tenuis, marginibus elevatis, pagina hirsuta. Hymenocarpus resupinatus. Hymenium N reagens (roseus). Sporis elliptis, hyalinis, non septatis, 10–12 μm longis, 7–8 μm latis. Type:— VENEZUELA. Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua cloud forests, elev. 2650 m, 17 July 2021, V. Marcano 21-65 (holotype MER, isotype B, private herb. V. Marcano). Etymology:—The epithet is dedicated to the naturalist Laura Castillo who has contributed with the author to the study of lichens and in discovering new species. Description:— Thallus on liverworts or mosses, byssaceous, filamentous, forming a thin crust, epiphytic on tree branches. Thallus surface irregular, more or less continuous, covering up to 12 cm across, greenish blue, developing a mat of cyanobacterial fibrils leaving interspaces, bordered by a discontinuous, fibrillose, narrow, white margin projecting vertical from the substrate, 2–4 mm broad; surface hirsute. Thallus (when fresh) in section 1.75–2 mm thick, composed of a loose, white medulla (0.75–1 mm) (hypothallus) and a broad, very loose aeruginous upper layer (1–1.25 mm) composed of irregularly arranged, long, white fibrils mixed with photobiont filaments (Rhizonema); cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsaw puzzle-shaped cells; cyanobacterial filaments not divided, composed of 12–17 μm wide and 7–8 μm high, bluish green cells penetrated by tubular fungal hyphae (haustoria); heterocytes frequent, bright yellow, elliptic, 10–12 μm wide and 5–8 μm high; hyphae associated with hyphal sheath straight, hyaline, 5–7 μm thick, smooth towards the tips of the filaments; clamp connections absent; hyphal sheath waxy in lateral outline. Hymenophore (when fresh) developed as small, angular to elongate, white, resupinate patches dispersed on the underside and resembling small, isomorphic attachment, small hapters (250–300 × 300–500 μm) with white, finely reticulate surface and entire margins; hymenophore composed of a paraplectenchymatous layer connected to loose medullary hyphae, 6.25–7.5 μm wide; hymenium composed of numerous, palisade-like basidioles and scattered basidia; basidia 32–39 × 5–7 μm, 4-sterigmate; basidiospores elliptic, non-septate, hyaline, 10–12 × 7–8 μm. Chemistry:—Thallus K–, C–, KC–, P–; medulla K–, C–, KC–, P–, N + (pink), ER+ (pale bluish purple); basidiocarp K–, C–, KC–, P–, N –, ER– (n = 6). No lichen compounds detected by TLC. Distribution and Ecology:— Dictyonema laurae was found growing as an epiphyte on bryophytes in the canopy of partly shady and very humid rainforests at elev. 2450–2650 m (Fig. 8A). It is associated with Grammitis, Cladonia and Parmotrema species. Remarks:— Dictyonema laurae is characterized by a thallus forming a thin crust (1.75–2 mm), with an irregular, continuous, greenish blue, hirsute surface, with discontinuous, fibrillose, narrow margins ascending from the substrate, hapteriform hymenophore patches with finely reticulate surface and entire margins, absence of clamp connections on the hyphae, large basidiospores, and a positive N spot test (pink). Dictyonema laurae is similar to D. subsericeum; which occupies the same habitat and has hymenophores resembling attachment hapters. However, D. subsericeum has a thallus forming a thicker crust (3–5 mm), with filamentous lobes, semicircular or hemispherical, projecting horizontally from the substrate, is variable in color (white, greenish blue, brown), size and shape (semicircular and hemispherical), with clamp connections on the hyphae, wider cyanobacterial cells, with a larger, irregular hapteriform hymenophore with more or less smooth or pruinose surface and fibrillose margins, smaller basidiospores, and a negative N spot test. Dictyonema laurae is also like D. hapteriferum Lücking, Dal Forno & Wilk (Lücking et al. 2013b: 11) and D. subinvolutum. However, D. hapteriferum is characterized by an absence of clamp connections, cyanobacterial filaments composed of 10–12 μm wide cells, a hymenophore resembling large attachment hapters but has semicircular, shelf-like, filamentous lobes and hymenophore with beige surface; it is known only from Bolivia and Perú. Dictyonema subinvolutum has a thallus developing a mat of interwoven fungal-cyanobacterial fibrils, leaving interspaces connected by a defined hypothallus; the cyanobacterial filaments are 12–15 μm wide, have clamp connections, and the basidiocarp surface is pruinose with slightly involute margins. Both species D. laurae and D. subinvolutum show a positive N spot test (pink). Additional specimens examined (paratypes):— Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua cloud forests, elev. 2450 m, 17 July 2021, V. Marcano 21-78, 21-79 (MER, private herb. V. Marcano)., Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on pages 213-215, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966, {"references":["Lucking, R., Dal Forno, M., Wilk, K. & Lawrey, J. D. (2013 b) Three new species of Dictyonema (lichenized Basidiomycota: Hygrophoraceae) from Bolivia. Acta Nova 6: 4 - 16."]}

2) Cyphellostereum mucuyense V. Marcano , sp. nov.; (Fig. 5) Mycobank MB#838446 Diagnosis:— Sicut Dictyonema metallicum, sed cellulis quadrato-rhombici nullis differt. Thallus muscicola vel terricola, sat tenuis. Prothallus nullus. Hyphae fibulatae. Haustoria nulla. Hymenocarpus resupinatus. Hymenium N non reagens. Basidiosporae subglobosae, 5–7 μm crassae. Type:— VENEZUELA. Mérida: Sierra Nevada de Mérida, San Jerónimo cloud forests, near Mucuy Alta, elev. 2150 m, terricolous; 30 October 2020, V. Marcano 20-567 (holotype MER, isotype B, private herb. V. Marcano). Etymology:—The epithet refers to the locality where the type specimens were collected. Description:— Thallus growing on and among bryophytes or soils, crustose-filamentous, dark green, developing a mat of ± individual to slightly interwoven fungal-cyanobacterial fibrils connected by a very thin hypothallus; prothallus absent. Fibrils horizontally arranged and closely appressed. Photobiont layer (50–120 μm) composed of cyanobacterial filaments covered by thin elongated fungal cells, not forming jigsaw-puzzle-shaped cells; cyanobacterial filaments not branched, composed of 8–10 μm wide and 4–9 μm high, green cells, mostly square to elongate–cylindrical, in chains, not penetrated by tubular fungal hyphae (haustoria); heterocytes rare, pale to slightly bright yellow, 7–8 μm wide and 4–5 μm high; cells of hyphal sheath regular in lateral outline, 2–4 μm thick; hyphae of medulla and hypothallus straight, hyaline, much branched, 3.5–5 μm thick. Clamp connections present, not common; hypothallus formed by interwoven, strongly agglutinate, generative hyphae. Hymenophore (when fresh) developed as effuse patches, resupinate-corticioid, 30–80 mm diameter, white, without distinct margins, with finely arachnoid surface; composed of a paraplectenchymatous layer resting on hyphae emerging from supporting thallus; in section 40–50 μm thick, composed of 3.5–5 μm thick hyphae; hymenium composed of numerous, palisade–like basidioles and scattered basidia; basidioles 20–30 × 5–6 μm; basidia 25–40 × 5–7 μm; basidiospores subglobose, non-septate, hyaline, 5–7 μm. Chemistry:—Thallus K–, C–, KC–, P–; medulla K–, C–, KC–, P–, N –, ER–; basidiocarp K–, C–, KC–, P–, N –, ER– (n = 6). No lichen compounds detected by TLC. Distribution and Ecology:—This species is known only from the well-developed type collection. It was found growing on mosses and acid soils in shady and very humid rainforest at 2150 m (Fig. 4A). This is the first record of the genus Cyphellostereum in Venezuela. Remarks:—Morphologically, C. mucuyense is a typical representative species of Cyphellostereum. It is characterized by an appressed filamentous thallus, a distinct, thin hypothallus, clamp connections, resupinate basidiomata (hymenophore) and absence of jigsaw-puzzle-shaped hyphal cells. Morphologically it resembles Dictyonema metallicum Lücking, Dal Forno & Lawrey (Lücking et al. 2013a: 22) in having appressed, horizontal fibrils and absence of a discernible medulla. The latter differs in the presence of a prothallus, an irregular photobiont layer, the dark greenish color of the cyanobacterial photobiont, a hyphal sheath formed by jigsaw-puzzle-shaped cells, and the absence of clamp connections. Cyphellostereum mucuyense could also be confused with D. obscuratum Lücking, Spielmann & Marcelli (Lücking et al. 2013a: 24) as both species are characterized by a strongly compressed mat of irregularly arranged, densely interwoven fibrils, but the latter species forms jigsaw-puzzle-shaped cells and lacks clamp connections (Lücking et al. 2013). Anatomically, C. mucuyense agrees with other Cyphellostereum species in the simple hyphal sheath around the cyanobacterial filaments, the comparatively narrow filament cells, and the absence of haustoria (Yanez et al. 2012, Lücking et al. 2013a, Dal Forno 2015; Dal Forno et al. 2013, 2016). The tiny fibrils of C. mucuyense are also reminiscent of those of Cyphellostereum. However, the resupinate-corticioid hymenophore is more reminiscent of Dictyonema s.str. (Lücking & Timdal 2016). Another species confirmed in the latter genus based on molecular data, D. huaorani, also lacks jigsaw-puzzle-shaped hyphal cells and haustoria (Schmull et al. 2014). Molecular studies are needed to determine the position of this taxon in the Dictyonema clade., Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on pages 208-210, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966, {"references":["Lucking, R., Dal Forno, M., Lawrey, J. D., Bungartz, F., Rojas, M. E. H., Hernandez M., J. E., Marcelli, M. P., Moncada, B., Morales, E. A., Nelsen, M. P., Paz, E., Salcedo, L., Spielmann, A. A., WilK, K., Will-Wolf, S. & Yanez-Ayabaca, A. (2013 a) Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales: Hygrophoraceae), with a key to all accepted genera and species in the Dictyonema clade. Phytotaxa 139: 1 - 38. https: // doi. org / 10.11646 / phytotaxa. 139.1.1","Yanez, A., Dal Forno, M., Bungartz, F., Lucking, R. & Lawrey, J. D. (2012) A first assessment of Galapagos basidiolichens. Fungal Diversity 52: 225 - 244. https: // doi. org / 10.1639 / 0007 - 2745 - 117.4.368","Dal Forno, M. (2015) Evolution and Diversity of the basidiolichen clade Dictyonema (Agaricales: Hygrophoraceae). PhD Dissertation, pp. 1 - 294.","Dal Forno, M., Lawrey, J. D., Sikaroodi, M., Bhattarai, S., Gillevet, P. M., Sulzbacher, M. & Lucking, R. (2013) Starting from scratch: evolution of the lichen thallus in the basidiolichen Dictyonema (Agaricales: Hygrophoraceae). Fungal Biology 117: 584 - 598. https: // doi. org / 10.1016 / j. funbio. 2013.05.006","Dal Forno, M., Lucking, R., Bungartz, F., Yanez-Ayabaca, A., Marcelli, M., Speilmann, A. A., Coca, L. F., Chaves, J. L., Aptroot, A., Sipman, H. J. M., Gillevet, P. & Lawrey, J. D. (2016) From one six: Unrecognized species diversity in the genus Acantholichen (lichenized Basidiomycota: Hygrophoraceae). Mycologia 108 (1): 38 - 55. https: // doi. org / 10.3852 / 15 - 060","Lucking, R. & Timdal, E. (2016) New species of Dictyonema and Cyphellostereum (lichenized Basidiomycota: Hygrophoraceae) from tropical Africa and the Indian Ocean, dedicated to the late Hildur Krog. Willdenowia 46: 191 - 199. https: // doi. org / 10.3372 / wi. 46.46115","Schmull, M., Dal Forno, M., Lucking, R., Cao, S., Clardy, J. & Lawrey, J. D. (2014) Dictyonema huaorani (Agaricales: Hygrophoraceae), a new lichenized basidiomycete from Amazonian Ecuador with presumed hallucinogenic properties. The Bryologist 117: 386 - 394. https: // doi. org / 10.1639 / 0007 - 2745 - 117.4.386"]}

3) Dictyonema andinum V. Marcano , sp. nov.; (Fig. 6) Mycobank MB#838341 Diagnosis:— Sicut Dictyonema irrigatum, sed fibrilis ascendentibus vel verticalibus differt. Thallus muscicola vel terricola, crassiusculus. Hyphis tegentibus fibrillas quadrato-rhombicis, undulatis, fibulatis. Haustoria praesentia. Filamenta 20–25 μm lata. Hymenocarpus corticoideus. Hymenium N reagens (rubescens), sporis guttaeformibus, hyalinis, non septatis, 5–6 μm longis. Type:— VENEZUELA. Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua, cloud forests, elev. 2650 m; 12 October 2020, V. Marcano & L. Castillo no. 20-535 (holotype MER, isotype B, private herb. V. Marcano). Etymology:—The epithet refers to the geographical region where the type specimens were collected. Description:— Thallus filamentous, dark blue-green, covering larger areas of the substrate, in irregular, confluent patches up to 45 cm across, forming a crust usually over bryophytes or soil, composed of loosely interwoven, dark green to bluish green fibrils leaving interspaces and bordered by a discrete, white margin (prothallus). Fibrils irregularly arranged, ascending or erect, forming an irregular, lumpy surface. Thallus furnished with irregular, small tufts, projecting vertically from the substrate, consisting of pale brown or green fibrils. Thallus (when fresh) in section 2–2.4 mm thick, composed of a thick photobiont (Rhizonema) layer (1.25–1.75 mm) and a thin, pale brown medulla (375–500 μm) (hypothallus); photobiont layer composed of numerous cyanobacterial filaments wrapped in a closed hyphal sheath formed by jigsaw-puzzle-shaped cells; cyanobacterial filaments not branched, composed of 20–25 μm wide and 7–8(–10) μm high, bluish green cells penetrated by tubular fungal hyphae (haustoria); heterocytes frequent, pale to slightly bright yellow, 10–12(–14) μm wide and 3–4(–5) μm high; cells of hyphal sheath variably wavy in lateral outline, 4–6 μm thick; hyphae of medulla, hypothallus and white margin (prothallus) straight, hyaline, much branched, 5–7 μm thick. Clamp connections present but sparse on the hyphae; hyphal sheath finely papillose towards the tips of the filaments; hypothallus and prothallus formed by interwoven, strongly agglutinate, generative hyphae; often fertile. Hymenophore (when fresh) developed as bulging, stereoid-corticioid patches from the underside of the thallus margins and soon becoming inverted and exposed, white patches, 6–14 mm wide; smooth, with pruinose upper surface and spongy lower surface; hymenophore in section 150–350 μm thick, composed of a paraplectenchymatous layer connected to loose medullary hyphae; hymenium composed of numerous, palisade-like basidioles and scattered basidia; basidioles 25–35 × 5–6 μm; basidia 30–40 × 5–7 μm, 4–sterigmate; basidiospores ellipsoid to narrowly drop-shaped, non-septate, hyaline, 5–6(–7) × 3.8–4(–5) μm. Chemistry:—Thallus K–, C–, KC–, P–; medulla K–, C–, KC–, P–, N + (dark red), ER+ (purple); basidiocarp K–, C–, KC–, P–, N + (red), ER+ (bluish purple) (n = 8). No lichen compounds detected by TLC. Distribution and Ecology:— D. andinum is the most abundant of all the species of Dictyonema s.str. found in the Venezuelan Andes. Six populations were observed in an area ca. 2 km 2. It was found growing on mosses and acid soils, associated with Dibaeis absoluta and Cladonia ceratophylla (Swartz 1788: 147) Sprengel (1827: 271) in partly exposed and very humid rainforest at 2000–2450 m (Fig. 5A). Remarks:—Morphologically, D. andinum is a typical representative of Dictyonema s.str. This species is characterized by irregularly arranged, broad, ascending, or erect fibrils, thick photobiont layer and hypothallus, finely papillose hyphae towards the tips of the filaments, clamp connections, and a white, stereoid-corticioid hymenophore. Commonly the thallus is slightly thicker when the specimen is fresh (humid). Positive (red) N spot test in the hymenophore and medulla are notable and suggest the presence of certain compounds which are absent in other Dictyonema species. Dictyonema andinum could be confused with D. irrigatum (Berkeley & M.A. Curtis 1860: 123) Lücking (2013b: 30). However, D. irrigatum has irregularly appressed, horizontal to erect fibrils, and a hymenophore developed as resupinate patches. Its spores are also slightly longer (7–9 μm) than those of D. andinum. Dictyonema irrigatum is known only from China and the Andaman and Nicobar Islands (Lücking et al. 2013 a, Jagadesh-Ram & Singh 2014) (Table 1). Additional specimens examined (paratypes):— Mérida: Parque Sierra Nevada de Mérida, Raiz de Agua, cloud forests, elev. 2650 m; 12 October 2020, V. Marcano & L. Castillo no. 20-540 (MER, private herb. V. Marcano); Parque Sierra Nevada de Mérida, road towards Lake Coromoto, cloud forest, elev. 2000 m, V. Marcano & L. Castillo 21-119, 21-120 (MER, private herb. V. Marcano)., Published as part of Marcano, Vicente, 2022, Eight new species of lichenized Basidiomycota in the genera Acantholichen, Cyphellostereum and Dictyonema s. str. (Agaricales, Hygrophoraceae) from northern South America, pp. 199-225 in Phytotaxa 574 (3) on pages 210-211, DOI: 10.11646/phytotaxa.574.3.1, http://zenodo.org/record/7380966, {"references":["Swartz, O. P. (1788) Nova Genera et Species Plantarum seu Prodromus descriptionum Vegetabilium, maximam partern incognitorum quae sub itinere in Indiam Occidentalem annis 1783 - 87. Holmiae in bibliopolis academie Swideri, Stockholm, Uppsala, pp. 1 - 152. https: // doi. org / 10.5962 / bhl. title. 433","Sprengel, C. P. J. (1827) Systema Vegetabilium. Editio decima sexta / curante Curtio Sprengel, equite stellae Polaris et aquilae rubrae, volume 4, Pars I, Carl von Linne, Bibliotheque de Palais des Arts, pp. 1 - 592.","Lucking, R., Dal Forno, M., Wilk, K. & Lawrey, J. D. (2013 b) Three new species of Dictyonema (lichenized Basidiomycota: Hygrophoraceae) from Bolivia. Acta Nova 6: 4 - 16.","Lucking, R., Dal Forno, M., Lawrey, J. D., Bungartz, F., Rojas, M. E. H., Hernandez M., J. E., Marcelli, M. P., Moncada, B., Morales, E. A., Nelsen, M. P., Paz, E., Salcedo, L., Spielmann, A. A., WilK, K., Will-Wolf, S. & Yanez-Ayabaca, A. (2013 a) Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales: Hygrophoraceae), with a key to all accepted genera and species in the Dictyonema clade. Phytotaxa 139: 1 - 38. https: // doi. org / 10.11646 / phytotaxa. 139.1.1","Jagadesh-Ram, T. A. M. & Singh, K. P. (2014) Dictyonema irrigatum (Berkeley & M. A. Curtis) Lucking, an interesting basidiolichen from Great Nicobar Island, India. Indian Journal of Forestry 37 (3): 361 - 364. https: // doi. org / 10.54207 / bsmps 1000 - 2014 - XEB 66 K"]}

The Chinese species complex of Hygrophorus russula in subsection Clitocyboides has been studied. Three new species, H. orientalis, H. qinggangjun and H. yunnanensis were proposed, based on morphology, molecular systematics and chemical reactions. Hygrophorus qinggangjun and H. yunnanensis occur at high elevations of Yunnan China; H. orientalis is widely distributed in East Asia and occurs at low elevations. At present, seven taxa have been revealed in the H. russula species complex; a key to these species is provided.

Bellanger, Jean-Michel, Moreau, Pierre-Arthur, Dima, Bálint, Albert, László, Krisai-Greilhuber, Irmgard, Campo, Emanuele, Sleiman, Sandra, Dvořák, Daniel, Zehnálek, Petr, Holec, Jan (2022): Hygrophorus roseodiscoideus Bon & Chevassut: epitypification and updated biogeography of a poorly known but widespread thermophilous species. Cryptogamie, Mycologie 20 (4): 81-90, DOI: 10.5252/cryptogamie-mycologie2022v43a4, URL: http://dx.doi.org/10.5252/cryptogamie-mycologie2022v43a4

Hygrophorus roseodiscoideus Bon & Chevassut Documents mycologiques 15 (59): 42 (Bon & Chevassut 1985). ORIGINAL DIAGNOSIS. — Pileus 5-12 cm, convexus vel late mammosus, carnosus, margine involuta, cuticula viscosa, coloribus versus discum fulvobrunneis vel rubrocastaneis, versus marginem paulatim pallidioribus, aurantio-ochraceis vel cremeo-roseis. Lamellae arcuatae, decurrentes, distantes, pallide cremeo roseae, vel ab imo subincarnatae. Stipes 6-10(13) × 1-2.5 cm, deorsum attenuato, sursum albido-punctuato, zona viscosa subannuliformi, fugaci atque versus basim nonnulis fasciolis +/- ochraceis. Caro pallide incarnata, odore saporeque subnullo vel fungoso. Sporae 9-10 × 5.5-6.5 µm, ellipticae.Trama bilateralis; basidia 45-65 × 7-10 µm, tetraspora. Cuticula pilei hyphis × 3-5 µm, gelatinosis, pigmento mixto vel membranari interdum verrucoso. Caulocutis sursum pilis +/- refringentibus, deorsum hyphis elongatis in micaceo muco coercitis. In Querceto ilicis vel pubescentis lectus, serotinus. Holotypus no. 84 391 in herbario MB – Cotypus GC 3326. HOLOTYPE. — France. Gard, close to Quissac, Quercus ilex and Q. pubescens on calcareous bedrock, 12.XII.1984, leg. G. Chevassut, det. M. Bon, Bon 84391 (holo-, LIP[LIP 0102284]); iso-, in herb. G. Chevassut 3326 (current location of this herbarium is unknown) (Fig. 1). EPITYPE. — France. Bouches-du-Rhône, Simiane-Collongue, calcareous bedrock, under Q. ilex and Pinus halepensis Mill., 31.X.2000, leg. & det. P.-A. Moreau, PAM00103101 (epi-, designated here in support of the holotype cited above, LIP[LIP 0402283]), GenBank ITS: MZ576440, MycoBank: MBT 10003653 (Fig. 2). ADDITIONAL MATERIAL EXAMINED. — Austria. Burgenland, district Oberpullendorf, community Horitschon, Ragerwald, 47°34’30”N, 16°32’30”E, 290 m a.s.l., bedrock, Inner Alpine Neogene Basin, clay, marl, marlstone, sand, gravel, limestone, in thermophilous oak forest, near Quercus cerris, Carpinus betulus L., and Acer L., c. 10 basidiomata, 29.IX.1988, leg. W. Klofac, det. A. Hausknecht (WU[WU-Myc 007470]). Croatia. Istria Region, Novigrad, Tar, sea coast, Mediterranean area, under Quercus ilex and Arbutus unedo L., 6.XII.2003, leg. & det. E. Campo (TUR [TUR-A 162859]); under Quercus pubescens, on calcarous soil, 6.XII.2014, leg.& det. E. Campo (MCVE [MCVE 28439]); Kvarner Region, Krk, Malinska, in Mediterranean area, on calcareous soil under Q. pubescens and Q. cerris, 17.XI.2018, leg. & det. E. Campo (specimens not kept). Czech Republic. Central Bohemia, near the village of Mcely, Čtvrtě National Nature Reserve, 50°17’54”N, 15°4’11.579”E, 225 m a.s.l.,under Carpinus L., Tilia L., Corylus L. and Quercus robur L. on calcareous bedrock (arenaceous marl), 10.IX.2010, leg. Jan Borovička, voucher lost; 50°17’39.96”N, 15°3’9.84”E, 250 m a.s.l., thermophilous oak forest, calcareous bedrock, under Quercus robur, 5.XI.2020, leg. & det. J. Holec, JH366/2020 (PRM [PRM 955166]; Fig. 4A); 50°17’52.868”N, 15°4’15.354”E, 225 m a.s.l., in warm broadleaved forest under Carpinus and Quercus, 1.XI.2020, leg. & det. J. Borovička, 3192 (PRM [PRM 954665]) (This site is 1.4 km NEE of the previous one and very close [about 50-100 m] to the first record from Čtvrtě in September 2010 [see above].); South Moravia, near Mokrá-Horákov, Sivický les, 49°13’16.2”N, 16°45’46.439”E (accuracy several hundred meters), 350-400 m a.s.l., thermophilous oak forest, calcareous bedrock, under Quercus petraea (Matt.) Liebl., Carpinus and Sorbus torminalis (L.) Crantz, 4.XI.2007, leg. & det. D. Dvořák, DD 501/07 (BRNU [BRNU 677820]; Fig. 4B). France. Bouches-du-Rhône, Simiane-Collongue, calcareous bedrock, under Q. ilex and Pinus halepensis, 31.X.2000, leg.& det.P.-A.Moreau, PAM00103101 (epi-, LIP[LIP 0402283]; Fig. 2); Hérault, Montpellier, campus du CEFE-CNRS, calcareous bedrock, under Q. ilex and Pinus halepensis, 5.XII.2010, leg. & det. F. Richard, FR0512001 (CEFE); Hérault, Puechabon, site expérimental du CEFE-CNRS, 43°44’29”N, 03°35’45”E, 270 m a.s.l., calcareous bedrock, soil under Q. ilex, 28.I.2009, leg. O. Shahin, det. F. Richard (environmental sample VAC25b-09), GenBank: JF506763 (Richard et al. 2011); Vaucluse, Cadenet, La Royère, calcareous bedrock, under Q. ilex, Q. pubescens and P. halepensis, 7.XII.2004, leg. & det. D. Borgarino, DiBo041144; Vaucluse, Cucuron, La Capagière, calcareous bedrock, under Q. ilex, Q. pubescens and P. halepensis, 10.XII.2004, leg. & det. D. Borgarino, DiBo041202 (Fig. 4C). Hungary. Budai-hegység Mts., Budapest, Mt. Széchenyi-hegy, 47°29’16.56”N, 18°58’56.279”E, 400 m a.s.l., thermophilous oak forest (Ceraso mahaleb -Quercetum pubescentis), calcareous bedrock, under Quercus pubescens, Quercus cerris, Fraxinus ornus L., Tilia cordata Mill., Prunus mahaleb L., Cornus mas L., 8.XI.2015, leg. & det. L. Albert, AL 280/15; 12.XI.2020, leg. & det. L. Albert & B. Dima, AL 342/20, DB 2020-11-12-2 (ELTE) (Fig. 4D); Budai-hegység Mts., Budapest, Mt. János-hegy, 47°31’18”N, 18°56’59.7”E, thermophilous oak forest, calcareous bedrock, under Quercus pubescens, Quercus cerris, Fraxinus ornus, 10.X.2014, leg. & det. B. Dima, DB5586 (ELTE); Budai-hegység Mts., Budakeszi, Wildlife Park, in deciduous forest, calcareous bedrock, under Quercus petraea, Carpinus betulus, 21.X.1976, leg. G. Bohus & M. Babos (as Hygrophorus fagi G.Becker & Bon), det. B. Dima (BP [BP58973]), GenBank: MK088109 (Zajta et al. 2019); Bükk Mts., Borsod-Abaúj-Zemplén County, Miskolc, Csanyik-völgy, in deciduous forest, calcareous bedrock, under Carpinus betulus, Fagus sylvatica L., Quercus petraea, 16.X.2007, leg. & det. L. Albert & B. Dima, AL 07/90; Vértes Mts., Fejér County, Csákvár, thermophilous oak forest, calcareous bedrock, under Quercus cerris, 1.XI.2015, leg. P. Finy, det. B. Dima, FP 2015-11-01. Italy. Friuli Venezia Giulia,Trieste, Parco di Miramare, 45°42’16.18”N, 13°42’52.37”E, 10 m a.s.l., on calcareous soil under Q. ilex, 28.XII.2012, leg. & det. P. Picciola (AMB-Muggia[AMB-Muggia 5824]); 21.XII.2014 (AMB-Muggia[AMB-Muggia 5657]); 28.XI.2015 (AMB-Muggia[AMB-Muggia 5846]); Friuli Venezia Giulia,Trieste, Muggia, Santa Barbara, 45°35’14.89”N, 13°47’4.50”E, 200 m a.s.l., on calcareous soil in mixed wood with Q. pubescens and P. nigra J.F.Arnold, 11.X.2017, leg. & det. P. Picciola (AMB-Muggia[AMB-Muggia 7196]); Friuli Venezia Giulia, Trieste, Basovizza, Bosco Igouza, 45°39’14.47”N, 13°52’24.48”E, 400 m a.s.l., on calcareous soil in mexed wood with P. nigra, Quercus sp. and other deciduous trees, 15.X.2017, leg. & det. P. Picciola (AMB-Muggia[AMB-Muggia 7175]); Toscana, Siena, Castellina in Chianti, Cipresseta S. Agnese, on calcareous soil under Quercus pubescens and Q. ilex, 28.X.2016, leg. & det. E. Campo, EC16102825 (Fig. 4E); Umbria, Carpegna, Passo Cantoniera, deciduous forest with Quercus cerris, 13.X.2004, leg. & det. A. Hausknecht (WU [WU-Myc 25200]; Fig. 4F). Lebanon. North Lebanon, Akkar district, near the village of Tesheh, 34°29’24.048”N, 36°11’34.343”E, 1354 m a.s.l., mixed limestone, sandstone and volcanic soil in Quercus calliprinos Webb forest with understory of Styrax officinalis L. and Juniperus oxycedrus L., 15.XII.2018, leg. & det. S. Sleiman (CEFE), GenBank:MZ088125 (Sleiman et al. 2021). ECOLOGY AND DISTRIBUTION. — Austria. The only known locality (Burgenland, Ragerwald) is a thermophilous oak forest with Quercus cerris on calcareous bedrock. Alltogether, some 500 species of fungi are known from this forest, among them rare in northern localities thermophilic species like Amanita caesarea (Scop.) Pers., Neoboletus xanthopus (Klofac & A.Urb.) Klofac & A.Urb., Rubroboletus legaliae (Pilát & Dermek) Della Maggiora & Trassinelli, R. rhodoxanthus (Krombh.) Kuan Zhao & Zhu L. Yang, R. satanas (Lenz) Kuan Zhao & Zhu L. Yang, Russula pannonica Pidlich-Aigner, as well as Omphalotus olearius (DC.) Singer, Desarmillaria tabescens (Scop.) R.A.Koch & Aime, Butyriboletus regius (Krombh.) D.Arora & J.L.Frank, and Astraeus hygrometricus (Pers.) Morgan. Croatia. The only records known to us are from the coastal Mediterranean areas of North Croatia (Istria, Krk), on calcareous soils, under Quercus ilex, Q. pubescens or Q. cerris. Czech Republic. Hygrophorus roseodiscoideus is known only from two localities. Both of them are thermophilous oak forests on calcareous bedrock (arenaceous marl, limestone). In Čtvrtě (Central Bohemia), the forest is mainly composed by Quercus robur but also other trees like Q. pubescens, Carpinus betulus, Cornus mas, Sorbus torminalis, or Tilia. Interestingly, H. roseodiscoideus was not found at the hottest sites typical by occurrence of Q. pubescens, but all three collections were under Quercus robur. In Sivický les (Southern Moravia), the host tree was most likely Q. petraea. Čtvrtě site is characteristic by a typical community of thermophilous ectomycorrhizal fungi (Šebek 1990; Zíbarová 2014; Holec 2020) like Amanita solitaria (Paulet ex Vittad.) Bertill., A. franchetii (Boud.) Fayod, Butyriboletus appendiculatus (Schaeff.) D.Arora & J.L.Frank, B. fechtneri (Velen.) D.Arora & J.L.Frank, Entoloma sinuatum (Pers.) P.Kumm., Hemileccinum impolitum (Fr.) Šutara, Hygrophorus penarioides Jacobsson & E.Larss., Inocybe godeyi Gillet, Lactarius acerrimus Britzelm., L. evosmus Kühner & Romagn., L. pterosporus Romagn., Leccinellum crocipodium (Letellier) Della Maggiora & Trassinelli, L. pseudoscabrum (Kallenb.) Šutara, Russula decipiens (Singer) Svrček, R. luteotacta Rea and numerous Cortinarius (Pers.) Gray (Phlegmacium (Fr.) Wünsche) species. The locality is protected as a national nature reserve and about 500 species of fungi are recorded there. Sivický les hosts a rich community of mainly ectomycorrhizal thermophilous and calciphilous species (Antonín et al. 2007), e.g. Amanita solitaria, Aureoboletus gentilis (Quél.) Pouzar, Butyriboletus appendiculatus, Cortinarius cotoneus Fr., C. dibaphus Fr., C. sodagnitus Rob.Henry and other cortinarii, Hemileccinum depilatum (Redeuilh) Šutara, H. impolitum, Hygrophorus arbustivus, H. russula (Schaeff. ex Fr.) Kauffman, Lactarius zonarius (Bull.) Fr., Lyophyllum rhopalopodium Clémençon, Rubroboletus rhodoxanthus, R. satanas, Suillellus queletii (Schulzer) Vizzini, Simonini & Gelardi, Tricholoma filamentosum (Alessio) Alessio and many others. The site is currently declared as Special Area of Conservation (Natura 2000 site) with occurrence of Galio-Carpinetum oak-hornbeam forests (habitat 9170) and Euro-Siberian steppic woods with Quercus spp. (habitat 91I0). A small part of the area is protected as the Nature Monument Na Líchách. France. All molecularly confirmed collections of H. roseodiscoideus come from thermophilous, calciphilous Quercus (Q. ilex, Q. pubescens) woodlands of the Mediterranean area (cf. above). However, the species is reported from several localities in the western part of the country, under the influence of the Atlantic Ocean and always in the vicinity of Quercus trees, apparently following the geographical range of Q. ilex (cf. www.mycocharentes.fr and www.societemycologique-poitou.org). Although the identity of these collections remains to be confirmed, published photographs and descriptions are indeed compatible with H. roseodiscoideus and this species might be present there as well. Bon & Chevassut (1985) reported collections from acidic soils, but they appear to be less abundant than those from calcareous ground. Hungary. Hygrophorus roseodiscoideus is known from five localities, of which three are situated around the forests of the capital, Budapest. Two older collections were determined as Hygrophorus fagi (Babos 1989) and H. leporinus Fr. s. Moser (Albert 2007). All localities are thermophilous calcareous deciduous forests dominated by Quercus species. Two of the localities (Mt. Széchenyi-hegy and Mt. Jánoshegy) are dominated by Q. pubescens with additional tree and shrub species such as Q. cerris, Fraxinus ornus, Tilia cordata, Cornus mas, etc. These habitats are hot-spots for many rare and thermophilous/ Mediterranean macrofungi such as Cortinarius atrovirens subsp. ionochlorus (Maire) Vizzini & Gasparini, C. quercilicis (Chevassut & Rob. Henry) Melot, C. violaceipes Bidaud & Consiglio, C. flavoaurantians Boccardo, Clericuzio & Vizzini, C. diffractosuavis Chevassut & Rob. Henry, C. ochrolamellatus Ballarà et al., C. chevassutii Rob.Henry, C. prodigiosus B.Bušek et al., Hygrophorus russula, Tricholoma basirubens (Bon) A.Riva & Bon, Boletus aereus Bull. The three other localities are not so thermophilic, they are dominated by Q. cerris or Q. petraea with admixture of Carpinus betulus. Italy. Documented collections are from northern and central Italy (see section Additionnal material examined). In all cases, H. roseodiscoideus was collected on calcareous soil, mostly under Quercus ilex, but also under Q. pubescens or Q. cerris, sometimes in mixed tree communities with Pinus nigra. In addition, E. Campo observed the species in the regions of Veneto (commune Chioggia), Liguria (Portofino), Emilia Romagna (Bettola), Toscana (Castellina in Chianti, Lucca, Capalbio), Marche (Ancona), Lazio (Tolfa), Sardegna (Isili, Nurallao) and Abruzzo (Loreto Aprutino). In these regions the occurence was linked to Quercus (Q. ilex, Q. pubescens, Q. cerris) on calcareous soils. It is very curious that the species is not reported from southern Italy (G. Schirinzi, pers. comm.). Lebanon. The first Asian record of H. roseodiscoideus is from northern Lebanon, in Quercus calliprinos forest with Styrax officinalis and Juniperus oxycedrus understory. Other species found at this site the same day were Tricholoma argyraceum (Bull.) Gillet, Cortinarius caligatus Malençon, Cortinarius subbulliardioides Rob.Henry, Cortinarius violaceipes, Cortinarius puellaris Brandrud et al., Cortinarius luteocingulatus Bidaud & Fillion and Inocybe tenebrosa Quél. All of them are new records for Lebanon and notably, this ecosystem appears to be the richest in fungal diversity compared to Abies cilicica (Antoine & Kotschy) Carrière and Quercus cerris ecosystems in the region (Sleiman et al. 2021). NOTES In the protologue, Bon & Chevassut (1985) cited a number of collections from the Montpellier area and from Roussillon, in the current Occitanie Region, as well as from Quercus woodlands near Aix-en-Provence and further east along the Côte d’Azur in the Provence-Alpes-Côte d’Azur Region. In its original concept, H. roseodiscoideus is thus a species fruiting along the whole French Mediterranean coast and in nomenclatural terms, all this cited material can be considered as paratypic. Unfortunately, the holotype could not be successfully sequenced (neither by the “classical” Sanger method nor the Illumina Miseq method), and none of the paratypes cited by Bon & Chevassut (1985: 44) could be located in Bon’s collections at LIP. Georges Chevassut’s herbarium has been lost after his death and no collection from his herbarium could be located either. To fix usage of the name, we thus epitypify it here using a sequenced collection from Simiane-Collongue, a locality 15 km south from Aix-en-Provence. This collection perfectly fits the description of H. roseodiscoideus, both macro- and micro-anatomically (Fig. 2), and its ITS sequence can thus be used as a barcode for the species in future studies., Published as part of Bellanger, Jean-Michel, Moreau, Pierre-Arthur, Dima, Bálint, Albert, László, Krisai-Greilhuber, Irmgard, Campo, Emanuele, Sleiman, Sandra, Dvořák, Daniel, Zehnálek, Petr & Holec, Jan, 2022, Hygrophorus roseodiscoideus Bon & Chevassut: epitypification and updated biogeography of a poorly known but widespread thermophilous species, pp. 81-90 in Cryptogamie, Mycologie 20 (4) on pages 83-88, DOI: 10.5252/cryptogamie-mycologie2022v43a4, http://zenodo.org/record/7815305, {"references":["BON M. & CHEVASSUT G. 1985. - Trois Hygrophores nouveaux de la chenaie verte mediterraneenne et leur ecologie (Quercetum Gallo-provinciale). Documents mycologiques 15 (59): 41 - 50.","RICHARD F., ROY M., SHAHIN O., STHULTZ C., DUCHEMIN M., JOFFRE R. & SELOSSE M. - A. 2011. - Ectomycorrhizal communities in a Mediterranean forest ecosystem dominated by Quercus ilex: seasonal dynamics and response to drought in the surface organic horizon. Annals of Forest Science 68 (1): 57 - 68. https: // doi. org / 10.1007 / s 13595 - 010 - 0007 - 5","ZAJTA E., VARGA T., KOVACS G. M. & DIMA B. 2019. - New insights on the taxonomy and distribution of Hygrophorus penarioides and Hygrophorus penarius from Hungary. Phytotaxa 392 (2): 127 - 139. https: // doi. org / 10.11646 / phytotaxa. 392.2.2","SLEIMAN S., BELLANGER J. - M., RICHARD F. & STEPHAN J. 2021. - First molecular-based contribution to the checklist of Lebanon macrofungi. Mycotaxon 136 (3): 687 - 687. https: // doi. org / 10.5248 / 136.687","SEBEK S. 1990. - Prispevek k mykologicke charakteristice teplomilnych doubrav u Mcel (okr. Nymburk) (A contribution towards mycological characteristics of thermophilous oak forests near Mcely (district Nymburk )), in KUTHAN J. & KOTLABA F. (eds), Vyzkum a ochrana hub v prirodnich rezervacich. Czechoslovak Scientific Society for Mycology, Praha: 40 - 46. http: // www. czechmycology. org / _ pub / pub 16. pdf","ZIBAROVA L. 2014. - Zaverecna zprava z orientacniho mykologickeho pruzkumu NPR Ctvrte v l. 2012 a 2014 (Final report from mycological research of National Nature Reserve Ctvrte in the years 2012 and 2014). Research report, depon, in AOPK CR, Praha.","ANTONIN V., DVORAK D. & VAGNER A. 2007. - Makromycety EVL Sivicky les. Research report, depon., in AOPK CR, Praha and Administration of Moravian Karst LPA.","BABOS M. 1989. - Magyarorszag kalaposgombainak (Agaricales s. l.) jegyzeke (The Agaricales s. l. taxa of Hungary 1). Mikologiai Kozlemenyek, Clusiana 28 (1 - 3): 3 - 234.","ALBERT L. 2007. - Szines oldalak, colour pages: Hygrophorus leporinus s. Moser. Mikologiai Kozlemenyek, Clusiana 46 (2): 285 - 286."]}

Phylogenetically, the genera Cuphophyllus, Ampulloclitocybe and Cantharocybe are treated as basal in the family Hygrophoraceae, despite weak support. However, the exact phylogenetic positions of the three genera have remained unresolved, and taxa related to these genera are poorly known. In this study, a new clitocyboid genus Spodocybe was proposed based on multigenic phylogenetic inference datasets and morphological evidence. The analyses of ITS as well as two combined datasets ITS-nrLSU-rpb2 and ITS-nrLSU-rpb1-rpb2-tef1-α-atp6 supported that (1) Spodocybe formed a well-supported monophyletic clade; and (2) sisters Spodocybe and Ampulloclitocybe, along with Cantharocybe and Cuphophyllus also formed a monophyletic lineage, as sister to the rest of the Hygrophoraceae. Meanwhile, two new species, namely S. rugosiceps and S. bispora, from southwestern China, were documented and illustrated. These results support the new proposed genus Spodocybe, and that Spodocybe, Ampulloclitocybe, Cantharocybe and Cuphophyllus should be retained in the Hygrophoraceae as a new subfamily Cuphophylloideae.

The genus Hygroaster (Hygrophoraceae, Hygrophorineae) is a small genus with a tropical distribution. Based on morphological and phylogenetic analysis, two new species, H. madagascarensis and H. andasibensis, are described from Madagascar. They are the first species of this genus described from Africa. To stabilize the nomenclature, the type specimens of H. nodulisporus, the type species of the genus, and H. fucatus were studied. The initial combination of the name Hygrophorus nodulisporus in the genus Hygroaster was invalid and is thus validated here. In addition, our study revealed that H. fucatus belongs to the genus Asproinocybe in suborder Tricholomatineae, whose representatives also have nodulose basidiospores. A new combination A. fucata is proposed.

A popular edible species of Hygrophorus, locally named as Mitangjun or Qinggangjun, is widely distributed in southwestern China, and has long been named “Hygrophorus russula”. Morphologically, it is very similar to H. parvirussula and H. russula, but it can be distinguished from them by its usually dense pileus scales, distant, fragile, and pinkish white lamellae with pink to reddish discoloration of lamellar edge when mature, and white context when fresh and dried. Phylogenetically, this fungus forms an independent lineage according to the ITS analysis. Ecologically, it is typically associated with Quercus aquifolioides at high elevations (over 3000 m) in southwestern China. Therefore, it is treated as a new species, namely H. deliciosus.