Hebeloma anthracophilum (Photo: H. J. Beker)

Taxonomy

Types: FRANCE: Meurthe-et-Moselle, Foret de Haye, Nancy (48.6833°N, 6.1833°E, alt. approx. 225 m a.s.l.) on burnt soil in woodland, Sep. 1907, R. Maire (Lectotype. herbarium acc. no. MPU 1921, HJB1000098). Lectotype designated by Grilli, Maire J.-C., Moreau P.-A. Robich G. (Eds.), Compléments a la Flore des Chamignons Superieurs de Maroc de G. Malençon et R. Bertault. Confédération Européenne de Mycologie Méditerranéenne, Nice: 299-318. : (2009) page 299.

Heterotypic synonyms:

• Hebeloma birrus var. odoratulum Bohus, Annales Historico-Naturales Musei Nationalis Hungarici 83: 84 (1991)
• Hebeloma rubrofuscum Velen., Ceske Houby: 395 (1919) ["1920"]

Homotypic synonyms:

• Hebelomatis anthracophilum (Maire) Locq., Flore Mycologique Vol III - Text. Cortinariales A: 146 (1979) ["1977"]

• arrow_drop_downarrow_drop_upEtymology
  From anthracinus– coal-black, and philus– (Greek) loving. This emphasizes the firesites or burnt ground that this taxon favours.

• arrow_drop_downarrow_drop_upOriginal diagnosis
  Pileo campanulato vel hemisphaerico expanso, plus minusve umbonato, glabro, viscido, fulvo, fibrillis innatis griseis cinerascenti pruinoso, disco fuscescente 4–5 rarius 7 cm diam., margine primitus involuto, albo-tomentello, in adulto expanso floccoso. Stipite e farcto cavo, recto, cylindrico, albido fibrilloso sericeo nec non squamulis concoloribus exasperato, 5–8 x 0.5 rarius 1 cm. Cortina nulla. Lamellis confertis, ex albido argillaceis, demum ferrugineo-fuscis, sinuato-adnatis vel uncinatis, ventricosis, acie albida vix denticulata. Carne albida nec raphaniodora. Sporis in charta ferrugineo-fuscis, pruniformibus, verruculosis, 10–11 x 6 μm. Basidiis cylindraceo-claviformibus, 4 rarius 2 sporis; cystidiis nullis.

• arrow_drop_downarrow_drop_upEnglish translation
  Pileus campanulate to hemispherical, expanding, more or less umbonate, glabrous, viscid, red-brown, with innate grey fibrils, ash-grey pruinose, becoming brown at centre, 4–5 rarely 7 cm in diam., margin involute at first, whitish tomentose, in expanded mature state floccose. Stipe solid then hollow, straight, shiny white fibrillose, but certainly without a roughness of adpressed concolorous squamules, 5–8 × 0.5 rarely 1 cm. Cortina absent. Lamellae crowded, white then clay-coloured then reddish brown sordid brown, sinuate-adnate to uncinate, ventricose, with hardly denticulate edge. Context whitish without raphanoid smell. Spores in mass rusty brown, pip-shaped, verruculose, 10–11 × 6 μm. Basidia cylindrical to clavate, four- rarely two-spored. Cystidia absent.

Description

Description of Hebeloma anthracophilum based on 22 collections

• arrow_drop_downarrow_drop_upMacroscopic description
  Pileus: (20) 27–43 (50) mm diameter; shape often convex, occasionally umbonate or broadly umbonate, rarely applanate; characters occasionally remains of universal veil; margin characters often involute or smooth, rarely overhanging pileus or ribbed; viscosity tacky when moist; colour variation usually two color, rarely unicolour; colour at centre occasionally yellowish brown, rarely sepia, ochraceous, dark pinkish buff, umber or cinnamon.

  Lamellae: attachment emarginate, rarely adnate; maximum depth 3–7 mm; number of complete lamellae 44–70; presence of tears absent; white fimbriate edge often weak, occasionally present or absent.

  Cortina presence: no.

  Stipe: (15) 22–65 (80) x (3) 4–8 (12) {median} x (3) 5–9 (13) {basal} mm; stipe Q 2.9–17.4; base shape often clavate or cylindrical, rarely bulbous or subbulbous; floccosity often floccose, occasionally fibrillose, pruinose or pruinose at apex; rooting variable; thick rhizoids at base absent;

  Context: Texture firm; stipe interior often hollow or stuffed; stipe flesh discolouring variable rarely weak or very strongly; slenderness measure 3.7–24.9; smell often odourless, rarely cocoa, raphanoid or soap; taste often bitter, occasionally strongly bitter, weakly bitter or mild where recorded.

  Spore deposit colour: usually umber, occasionally sepia.

  Exsiccata characters: occasionally hard or pileus blackening.

• arrow_drop_downarrow_drop_upMicroscopic description
  Spores: shape amygdaloid, occasionally fusoid or limoniform, rarely navicular; colour in microscope often brown or yellow brown; guttules often no, rarely yes or weak. papilla often no, occasionally weak, rarely yes; Spore Code: O3 O4; (P2) P3; D3 D4.

  Basidia: 22–35 (37) x 6–8 μm; ave. Q 3.4–5.1; spore arrangement 4 spored;

  Cheilocystidia: main shape cylindrical, occasionally clavate, rarely gently clavate, clavate-lageniform or clavate-ventricose, tapering or ventricose; special features observed occasionally short, clamped septa or septa, rarely branching, many collapsed in exsiccata or mucronate; cheilocystidia ratios: A/M = 1.07–1.23; A/B = 0.94–1.24; B/M = 0.91–1.19.

  Pleurocystidia: usually none seen, rarely seen.

  Ixocutis: epicutis thickness (measured from exsiccata) up to 375 μm; ixocutis hyphae width up to 4 μm; ixocutis hyphae encrustation often yes, occasionally no; shape of trama elements beneath subcutis thickly sausage-shaped, occasionally ellipsoid up to 13 μm wide.

  Caulocystidia: Similar to cheilocystidia but larger, up to 100 μm.

• arrow_drop_downarrow_drop_upSpore measurements

• arrow_drop_downarrow_drop_upCheilocystidia measurements

• arrow_drop_downarrow_drop_upHabitat and distribution
  Hebeloma anthracophilum's preferred habitat appears to be woodland or deciduous woodland with burnt soil. Where only one possible associate was recorded, the most commonly recorded associate was Fagus (41.7%) but Castanea (41.7%) and Quercus (16.7%) were also recorded. In these cases the most commonly recorded family was Fagaceae (100.0%). We have additional records where Carpinus was recorded as a possible associate, but for these collections a number of possible associates were mentioned. Overall the most commonly recorded families are Fagaceae (94.4%) and Betulaceae (5.6%) The growth habit of our collections was usually scattered, occasionally gregarious and rarely caespitose.

  According to our current collections, the species is predominantly found in Europe (86.4%) but also found in Temperate Asia (13.6%). On these continents, collections has been found in the WWF biomes The World Wildlife Fund (WWF) have divided the world into 867 terrestrial ecoregions. The ecoregion here is estimated by mapping from the GPS coordinates of the collection using data made available by Dinerstein et al (2017). Use this webtool to explore the ecoregions visually or see a full list of current ecoregions on Wikipedia. temperate broadleaf & mixed forests (68.2%) and mediterranean forests, woodlands & scrub (27.3%), specifically including the ecoregions: Western European broadleaf forests (40.9%), Italian sclerophyllous and semi-deciduous forests (18.2%) and Crimean Submediterranean forest complex (13.6%). From collector information, it appears collections have been found only in the 1.4 Forest – Temperate IUCN habitat We map from the collector's description of the habitat to the International Union for Conservation of Nature (IUCN)'s definition using a standardised set of rules. Please see this page for a full list of IUCN habitats..

  Within Europe we have records from the Centre (Czech Republic, Switzerland, Belgium and Hungary), the Southeast (Italy), the Southwest (France) and the North (England). Specimens have been collected from 39.4°N to 53.7°N.

  Within Temperate Asia all our records are from Caucasus (Adygea).

• arrow_drop_downarrow_drop_upMolecular results
  Hebeloma anthracophilum is closely related with H. birrus. This is supported by almost all loci. The ITS is the only locus which does not support H. anthracophilum as monophyletic and distinct from H. birrus. In all other phylogenetic results the species is supported or even very well supported by bootstrap in addition to being monophyletic. We do have the ITS, V6 and V9 sequence of the type of H. birrus var. odoratulum and thus have molecular support for the synonymization of this variety of H. birrus with this clade that, based on morphological evidence, we here refer to as H. anthracophilum. We are not aware of any ITS sequences ascribable to H. anthracophilum or H. birrus from outside Europe.

• arrow_drop_downarrow_drop_upCommentary
  Hebeloma anthracophilum without annulus and with its tendency to root together with the short, mainly cylindrical cheilocystidia place this taxon firmly within H. sect. Scabrispora. However, when there is no obvious rooting it may be difficult, at least microscopically, to separate this taxon from some members of H. sect. Naviculospora. Within the group of taxa with primarily cylindrical cheilocystidia, H. anthracophilum may be distinguished by the spore size (11.0–11.9 × 5.4–6.7 μm) and the strongly and constantly loosening perispore. Maire described this taxon as exclusively growing on burnt ground and this appears to be correct. However, the fact that the ground has been burnt can sometimes be overlooked and the spore size and characters are more reliable parameters in determination. Many authors have in the past synonymized this taxon with H. birrus (see for example Vesterholt 2005). While most microscopic characters are similar for these two taxa, H. anthracophilum certainly has, on average, longer spores, at least 11 μm, while the spores of H. birrus are never as long as this. As noted above, molecularly these taxa can also be easily separated, but note that the separation is not seen with ITS sequences. There are also macroscopic characters by which these taxa can be separated, in particular Hebeloma anthracophilum tends to be smaller and more squat. We have synonymized both H. birrus var. odoratulum and H. rubrofuscum with H. anthracophilum. For the former we have molecular as well as morphological grounds for the synonymy; for the latter we have relied purely on morphology as the holotype of H. rubrofuscum was stored in a preserving liquid that, although not necessarily destroying the DNA, appears to have made it impossible to extract PCR-quality DNA from the existing material.

Geographic distribution

Phenology

• arrow_drop_downarrow_drop_upAdditional cited collections

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