Hebeloma crustuliniforme (Photo: H. J. Beker)

Taxonomy

Basionym:
Agaricus crustuliniformis Bull. ., Herb. Fr.: 308 (1787)

Types: Bulliard, Herb. Fr.: 128 (1787) tab. 308, lectotype (icon) designated by Vesterholt et al., Mycol. Prog. 13 (3): (2014) page 558 FRANCE: Côte-d'Or, La Combe de la Serpente (47.3024°N, 4.9463°E, alt. approx. 435 m a.s.l.) on calcareous soil and litter in coniferous woodland plantation under Cedrus libanotica ssp.atlantica, 11 Oct. 2010, H.J. Beker, M.L. Beker (Epitype. herbarium acc. no. BR MYCO 173989-68 (epitype), C C-F-90146 (isoepitype), HJB13713). Epitype designated by Vesterholt et al., Mycol. Prog. 13 (3): (2014) page 558 (MBT175887).

• arrow_drop_downarrow_drop_upType notes
  The lectotype (icon), selected by Vesterholt et al., was incorrectly referred to as the holotype but this was corrected under Art. 9.9. in Beker et al. 215 (2016).

Heterotypic synonyms:

• Hebeloma macrosporum Velen., Ceske Houby: 398 (1919) ["1920"]

Homotypic synonyms:

• Derminus crustuliniformis (Bull.) J. Schröt., Die Pilze Schlesiens 3: 583 (1889)
• Hebelomatis crustuliniformis (Bull.) Locq, Flore Mycologique Vol III - Text. Cortinariales A: 146 (1979) ["1977"]
• Hylophila crustuliniformis (Bull.) Quél., Enchiridion Fungorum in Europa Media et Praesertim in Gallia Vigentium: 99 (1886)
• Agaricus ossa J.F. Gmel., Caroli a Linné - Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis 2: 1404 (1792)

• arrow_drop_downarrow_drop_upEtymology
  From crustulum– small pastry and formis– in the form of. The species is in English often referred to as poison pie.

• arrow_drop_downarrow_drop_upOriginal diagnosis
  L’agaric Echaudé. Agaricus crustuliniformis ... Ce champignon est très commun en automne dans les bois, les prairies, sa superficie est unie, luisante très gluante dans les temps humides, dans l’état de jeunesse comme dans l’état de vieillesse il ressemble parfaitement tant par sa forme que par sa couleur a un échaudé, ses feuillets sont libres laissent un intervalle entre eux et le pédicule, il y a toujours pour un feuillet entier quatre feuillets qui ne le sont pas, le pédicule est toujours plus ou moins pelucheux … rien de plus curieux que la manière dont ce champignon est semé sur la terre, tantôt autour du pied d’un arbre mais a une distance de 8 ou 10 pieds, tantôt au milieu d’une prairie, à travers une forêt vous en rencontrer des peuplades prodigieuses formant des ronds très réguliers et plus ou moins étendus des sentiers tortueux de deux ou trois cents pieds de long, sur un pied ou environ de large espace occupé par 5 ou 6 champignons de la même espèce si rapprochés que vous en voyer mille sans apercevoir le pédicule d’un seul. N.B. la fig A [Fig. 20.4A (colour)] représente la coupe verticale de ce champignon … quelquefois on le trouve un ici un la dans les bois fréquentés il est alors fort difficile a reconnaitre car il n’a que des caractères vagues. Il est très âcre et très désagréable au goût.

• arrow_drop_downarrow_drop_upEnglish translation
  The Scalded Agaric. Agaricus crustuliniformis ... This mushroom is very common in autumn in the woods and meadows; its surface is smooth, shiny, very sticky in wet weather; both when young and when older, it perfectly resembles in form and in colour an echaudé [kind of biscuit that was dipped in hot water before being baked]; its lamellae are free leaving a gap between them and the stipe, for each complete lamella there are always four lamellulae; the stipe is always more or less floccose ... there is nothing more curious than this mushroom’s growth habit, sometimes around the base of a tree but at a distance of 8 or 10 feet, sometimes in the middle of a meadow; when crossing a forest one might come across prodigious tribes forming very regular rings stretching in winding paths for 200–300 feet, and in a space a foot or so wide there may be 5 or 6 mushrooms of the same species so close together that you can see a thousand mushrooms without noticing a single stipe. NB Fig A [Fig. 20.4A (colour)] is the vertical section of this mushroom ... sometimes it can be found here and there in a wood one frequents but it is very difficult to recognise because it has only vague characters. It is very acrid and unpleasant to taste.

Description

Description of Hebeloma crustuliniforme based on 114 collections

• arrow_drop_downarrow_drop_upMacroscopic description
  Pileus: (20) 37–91 (135) mm diameter; shape often convex, occasionally umbonate or broadly umbonate, rarely strongly umbonate or weakly umbonate; characters rarely remains of universal veil or spotting; margin characters often involute, occasionally smooth, rarely eroded or ribbed; viscosity tacky when moist; colour variation often unicolour, occasionally two color; colour at centre occasionally pale cream, cream or yellowish brown, rarely warm buff, dark pinkish buff, pale pinkish buff or ochraceous.

  Lamellae: attachment usually emarginate, rarely adnate, adnexed or free; maximum depth 3–9 mm; number of complete lamellae 60–100; presence of tears usually visible with naked eye, rarely absent or visible with x10 lens; white fimbriate edge often present, occasionally very strong, rarely weak.

  Cortina presence: no.

  Stipe: (25) 33–91 (115) x (6) 7–20 (26) {median} x (8) 11–21 (27) {basal} mm; stipe Q 1.7–11.5; base shape often clavate, occasionally cylindrical or bulbous, rarely subbulbous; floccosity floccose, rarely pruinose at apex; rooting usually no, rarely weak; thick rhizoids at base usually absent, occasionally present;

  Context: Texture firm; stipe interior often stuffed or hollow; stipe flesh discolouring often no, rarely yes or weak; slenderness measure 0.8–12.6; smell occasionally raphanoid, odourless or weakly raphanoid, rarely strongly raphanoid or farinaceous; taste often hot, mild or strongly raphanoid where recorded.

  Spore deposit colour: often brownish olive, occasionally yellowish brown.

  Exsiccata characters: rarely pale or shiny.

• arrow_drop_downarrow_drop_upMicroscopic description
  Spores: shape amygdaloid, rarely limoniform; colour in microscope occasionally yellow or yellow brown, rarely brown, brown pale, brown yellow or yellow pale; guttules usually yes, rarely no. papilla occasionally no or weak, rarely yes or very strongly; Spore Code: O2 O3; P0 P1 (P2); (D0) D1 (D2).

  Basidia: (20) 25–39 (40) x 6–10 μm; ave. Q 3.1–4.6; spore arrangement 4 spored;

  Cheilocystidia: main shape usually clavate-stipitate, often spathulate-stipitate, occasionally clavate-lageniform or clavate-ventricose or gently clavate, rarely capitate-stipitate, clavate or cylindrical; special features observed occasionally sinuate, median thickening or septa, rarely clamped septa, conglutinate, many collapsed in exsiccata or yellow contents; cheilocystidia ratios: A/M = 1.60–2.35; A/B = 1.52–2.32; B/M = 0.83–1.24.

  Pleurocystidia: none seen.

  Ixocutis: epicutis thickness (measured from exsiccata) up to 350 μm; ixocutis hyphae width up to 7 μm; ixocutis hyphae encrustation often yes, occasionally no; shape of trama elements beneath subcutis often cylindrical or thickly sausage-shaped, occasionally thinly sausage-shaped, rarely hyphae thick or spherical up to 24 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 crustuliniforme's preferred substrate appears to be grassy soil. Where only one possible associate was recorded, the most commonly recorded associate was Quercus (24.6%) but Pinus (23.1%), Tilia (12.3%), Populus (10.8%), Cedrus (7.7%), Arbutus (7.7%), Picea (3.1%), Fagus (3.1%), Betula (1.5%), Cistus (1.5%), Eucalyptus (1.5%), Gleditsia (1.5%) and Tsuga (1.5%) were also recorded. In these cases the most commonly recorded families were Pinaceae (36.4%), Fagaceae (27.3%), Malvaceae (12.1%), Salicaceae (10.6%) and Ericaceae (7.6%). We have additional records where Salix (2.3%), Pseudotsuga (2.3%), Carpinus (1.2%), Corylus (1.2%) and Abies (1.2%) were recorded as possible associates, but in these cases a number of possible associates were mentioned. Overall the most commonly recorded families are Pinaceae (46.5%), Fagaceae (38.4%), Salicaceae (11.6%), Malvaceae (9.3%) and Ericaceae (8.1%) The growth habit of our collections was occasionally scattered or gregarious and rarely solitary or caespitose.

  According to our current data, the species is found on multiple continents with collections found in Northern America (48.1%), Europe (41.4%), Temperate Asia (4.8%), Australasia (3.9%), Africa (1.0%) and Southern America (1.0%). 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 (45.7%), mediterranean forests, woodlands & scrub (33.3%) and temperate conifer forests (16.2%), specifically including the ecoregions: Atlantic coastal pine barrens (11.4%) and California coastal sage and chaparral (10.5%). From collector information, it appears collections have been found in the 1.4 Forest – Temperate (47.7%) and 14.5 Urban Areas (32.3%) IUCN habitats 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 Northern America we have records from Northeastern U.S.A. (New York, Pennsylvania, Ohio, New Jersey and Vermont), Southwestern U.S.A. (California), Eastern Canada (Quebec and Ontario), Northwestern U.S.A. (Washington and Oregon), Southeastern U.S.A. (Virginia) and Western Canada (British Columbia).

  Within Europe we have records from the Southwest (Spain, France, Channel Islands and Portugal), the Centre (Germany, Netherlands and Czech Republic), the North (England and Denmark) and the Southeast (Italy, North Macedonia and Bulgaria). Specimens have been collected from 39.4°N to 56.2°N.

  Within Temperate Asia all our records are from Western Asia (Cyprus).

  Within Australasia all our records are from Australia (Victoria).

  Within Africa all our records are from Macaronesia (Spain).

  Within Southern America all our records are from Southern South America (Chile).

• arrow_drop_downarrow_drop_upMolecular results
  Hebeloma crustuliniforme as defined here forms well supported clades with all loci tested, alone or concatenated. Hebeloma crustuliniforme is in molecular terms not a member of what we have referred to as the alpinum-complex, but in fact it is molecularly related to, although quite distinct from, H. aanenii, H. alpinum, H. geminatum and H. eburneum, the four species forming the core of the complex. It should be possible to identify the species by ITS.

• arrow_drop_downarrow_drop_upCommentary
  Hebeloma crustuliniforme posesses cystidia of the shape we consider typical for its subsection, H. subsect. Crustuliniformia. The species most likely corresponds to ICG5 of Aanen & Kuyper (1999). Hebeloma crustuliniforme can be separated from other species within H. subsect. Crustuliniformia based on the non arctic/alpine habitat, the number of complete lamellae that always exceeds 60, the larger spore size that is usually greater than 11 μm long and 6 μm wide and the narrower average apex width to the cheilocystidium, less than 8 μm. Further, the pileus is often rather more coloured (brown) than that of the other members of this complex. Hebeloma crustuliniforme is a member of the complex of species that have usually been determined as H. crustuliniforme. This complex also includes H. aanenii, H. geminatum, H. eburneum and H. alpinum.

Geographic distribution

Phenology

• arrow_drop_downarrow_drop_upAdditional cited collections

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