Hebeloma hiemale (Photo: H. J. Beker)

Taxonomy

• arrow_drop_downarrow_drop_upNomenclatural notes
  Hebeloma hiemale is often quoted as published in 1892. Volbracht (2006), however, indicates that up to page 46 and icones 106–150 were published in 1892, while pages 47–81 and icones 151–195 were published in 1898.

Types: Bresadola, Fung. Trident. 2 (11-13): 52 (1898) pl. 160.1, lectotype (icon) designated by Eberhardt et al., Fungal Biol. 120: (2015) ["2016"] page 98 (MBT202913) FRANCE: Normandy, Foret Andaines (48.5808°N, 0.4583°W, alt. approx. 170 m a.s.l.) on boggy soil and litter in mixed woodland under Fagus sp. and Quercus sp., 7 Oct. 2006, H.J. Beker (Epitype. herbarium acc. no. BR BR-MYCO 174905-14 (epitype), C C-F-92314 (isoepitype), HJB11704). Epitype designated by Eberhardt et al., Fungal Biol. 120: (2015) ["2016"] page 98 (MBT202768).

Heterotypic synonyms:

• Hebeloma oculatum Bruchet, Bull. Mens. Soc. Linn. Lyon 39, supplement 6: 126 (1970)
• Hebelomatis oculatum (Bruchet) Locq., Flore Mycologique Vol III - Text. Cortinariales A: 146 (1979) ["1977"]
• Gymnopilus permollis Murrill, Mycologia 4 (5): 252 (1912)
• Flammula permollis (Murrill) Murrill, Mycologia 4 (5): 262 (1912)

Homotypic synonyms:

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

• arrow_drop_downarrow_drop_upEtymology
  From hiemalis- winter and wintry, presumably to emphasise Bresadola's view that this species produces basiodomes in the autumn and during spring.

• arrow_drop_downarrow_drop_upOriginal diagnosis
  Pileus carnosulus, e convexo-subhemispherico explanato-gibbosus, vel depressus, viscosus, glaber, margine primitus involuto et albo-flocculoso, alutaceo-pallidus, centro crustulinus vel interdum zona lata, crustulina in medio pilei ornatus, 2–4.5 cm latus; lamellae confertae, ex albis argillaceo-subcinnarmomeae, acie albo-flocculosa, postice sinuato-adnatae vel adnexae et fere liberae; stipes albus, deorsum lutescens, e farcto subcavus, aequalis, subfibrillosus, apice albo-furfuraceus, 2–3 cm. longus, 5–7 cm. crassus; caro alba vix odora, sapore subamaro; sporae obverse piriformes, sub microscopio flavo-aureae, 12–13 x 6–7; basidia clavata 30–35 x 7–8. Sero autumno ad primum verum, gregaria in nemoribus, ad margines silvarum etc. Gocciadoro pro Tridentum, Val di Sole etc. Obs. Hebelomati crustuliniformi proxime accedit, a quo statura constanter minori, odore vix manifesto et tempore vegetationis satis videtur diversum.

• arrow_drop_downarrow_drop_upEnglish translation
  Pileus thin-fleshed, first convex-hemispherical then expanding with umbo or depressed, viscous, glabrous, with margin first involute and white flocculose, yellowish-pallid, at centre pinkish cinnamon, often in a broad zone, 2–4.5 cm broad; lamellae crowded, white then pinkish yellow to cinnamon, with white flocculose edge, sinuate-adnate or adnexed to almost free; stipe white, yellowish towards base, solid then hollow, equal, subfibrillose, apex white furfuraceous, 2–3 cm long, 5–7 mm broad; context white with hardly any odour, taste slightly bitter; spores obpyriform, yellow-golden under microscope, 12–13 × 6–7 μm; basidia clavate, 30–35 × 7–8 μm. Late autumn and spring, in groups in forest, at margin of forest, etc. Gocciadoro near Trento, Val di Sole etc. Observation: comes very close to Hebeloma crustuliniforme, differs sufficiently by the constantly smaller habit, less distinct odour, and time of producing basidiomes.

Description

Description of Hebeloma hiemale based on 325 collections

• arrow_drop_downarrow_drop_upMacroscopic description
  Pileus: (10) 14–50 (80) mm diameter; shape often convex, occasionally umbonate, rarely broadly umbonate, strongly umbonate or applanate; characters rarely spotting, hygrophanous, pruinose or pubescent; margin characters often smooth, occasionally involute, rarely eroded, crenulate, sulcate, ribbed, scalloped or serrate; viscosity tacky when moist; colour variation often unicolour or two color; colour at centre occasionally dark pinkish buff, yellowish brown or ochraceous, rarely clay-buff, cream, honey, warm buff, dark greyish buff, orange-brown, cinnamon, umber, clay-pink, greyish buff, fuscous, fawn, pale yellow or buff-yellow.

  Lamellae: attachment usually emarginate, rarely adnate, adnexed, decurrent or free; maximum depth 3–8 mm; number of complete lamellae 35–71; presence of tears occasionally absent, visible with naked eye or visible with x10 lens; white fimbriate edge usually present, occasionally weak, rarely absent.

  Cortina presence: no.

  Stipe: (13) 22–72 (122) x (2) 3–8 (14) {median} x (2) 3–10 (15) {basal} mm; stipe Q 2.5–17.5; base shape often cylindrical, occasionally clavate, rarely tapering, subbulbous or bulbous; floccosity occasionally pruinose, floccose or pruinose at apex, rarely velute, weakly floccose, fibrillose, floccose at apex, none or fibrous; rooting usually no, rarely weak; thick rhizoids at base usually absent, rarely present;

  Context: Texture firm; stipe interior often hollow, occasionally stuffed, rarely superior wick or basal wick; stipe flesh discolouring often no, occasionally weak or yes, rarely very strongly; slenderness measure 1.4–35.8; smell often raphanoid, rarely odourless, weakly raphanoid, cocoa, strongly raphanoid, fruit or soap; taste occasionally mild, bitter, weakly bitter, hot or raphanoid, rarely strongly raphanoid or weakly raphanoid where recorded.

  Spore deposit colour: often brownish olive, occasionally yellowish brown, rarely umber, greyish brown or cinnamon.

  Exsiccata characters: occasionally pale, rarely lamellae blackening, dark, fragile, twisted stipe or pileus blackening.

• arrow_drop_downarrow_drop_upMicroscopic description
  Spores: shape usually amygdaloid, rarely limoniform, fusoid or ovoid; colour in microscope occasionally yellow brown, yellow or brown, rarely brown yellow or grey yellow; guttules often yes, occasionally no, rarely weak. papilla often yes, occasionally weak, rarely no; Spore Code: O2 O3; P0 P1; D1 D2.

  Basidia: (20) 23–38 (43) x (5) 6–10 μm; ave. Q 3.2–4.4; spore arrangement 4 spored;

  Cheilocystidia: main shape usually clavate-lageniform or clavate-ventricose, often clavate-stipitate, rarely clavate, gently clavate, ventricose, capitate-stipitate, lageniform, spathulate-stipitate, tapering or cylindrical; special features observed often median thickening, occasionally septa, apical thickening or geniculate, rarely spathulate, many collapsed in exsiccata, sinuate, slender, bifurcate, short, yellow contents, large, mucous, rostrate or uniform; cheilocystidia ratios: A/M = 1.45–2.51; A/B = 0.91–1.80; B/M = 1.26–1.61.

  Pleurocystidia: none seen.

  Ixocutis: epicutis thickness (measured from exsiccata) up to 160 μm; ixocutis hyphae width up to 7 μm; ixocutis hyphae encrustation usually yes, rarely no; shape of trama elements beneath subcutis occasionally cylindrical, isodiametric, angular or thickly sausage-shaped, rarely ellipsoid, thinly sausage-shaped or undefined up to 17 μm wide.

  Caulocystidia: Similar to cheilocystidia but larger, up to 105 μm.

• arrow_drop_downarrow_drop_upSpore measurements

• arrow_drop_downarrow_drop_upCheilocystidia measurements

• arrow_drop_downarrow_drop_upHabitat and distribution
  Where only one possible associate was recorded, the most commonly recorded associate was Salix (42.8%) but Betula (15.6%), Picea (9.2%), Pinus (6.4%), Dryas (5.8%), Quercus (5.2%), Fagus (4.6%), Populus (4.0%), Tilia (2.3%), Carpinus (1.2%) and Corylus (1.2%) were also recorded. In these cases the most commonly recorded families were Salicaceae (42.1%), Pinaceae (19.7%), Betulaceae (19.1%) and Fagaceae (10.9%). We have additional records where Alnus (4.8%), Abies (2.4%), Larix (1.7%) and Tsuga (1.4%) were recorded as possible associates, but in these cases a number of possible associates were mentioned. Overall the most commonly recorded families are Salicaceae (51.9%), Betulaceae (33.6%), Pinaceae (23.5%), Fagaceae (15.2%) and Rosaceae (6.6%) The growth habit of our collections was often scattered, occasionally solitary and rarely gregarious or caespitose.

  According to our current data, the species is found on multiple continents with collections found in Europe (59.1%), Northern America (38.1%), Temperate Asia (1.6%) and Australasia (1.2%). 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 (40.4%), temperate conifer forests (16.1%), tundra (15.8%), unknown biome (13.0%) and boreal forests/taiga (10.2%), specifically including the ecoregions: Unknown region (13.0%). From collector information, it appears collections have been found in the 1.4 Forest – Temperate (39.4%) and 4.1 Grassland – Tundra (21.5%) 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 Europe we have records from the North (Norway, England, Isle Of Man, Denmark, Scotland, Svalbard, Iceland, Finland, Sweden and Faroe Islands), the Centre (Belgium, Germany, Poland, Czech Republic and Netherlands), the Southwest (Spain and France), Eastern Europe (Estonia) and the Southeast (Italy). Specimens have been collected from 36.6°N to 78.9°N.

  Within Northern America we have records from Subarctic America (Greenland, Northwest Territories, Alaska and Nunavut), Northwestern U.S.A. (Colorado, Washington, Oregon, Montana and Wyoming), Eastern Canada (Newfoundland and Labrador and Quebec), Western Canada (British Columbia and Alberta), North-central U.S.A. (Minnesota and Wisconsin) and Southwestern U.S.A. (California and Arizona).

  Within Temperate Asia we have records from Caucasus (Karachay-Balkar), Russian Far East (Magadan Oblast) and Siberia (Tyumen).

  Within Australasia we have records from New Zealand (New Zealand) and Australia (Tasmania).

• arrow_drop_downarrow_drop_upMolecular results
  Beker et al. (2010) noted the rather high intraspecific and intragenomic variation within the ITS of H. hiemale. This is also true for other loci tested. However, we have not yet found any indication of cryptic species here.

• arrow_drop_downarrow_drop_upCommentary
  The main morphological features that distinguish species of H. subsect. Hiemalia are the shape of the cheilocystidium, significantly swollen at the apex and also swollen in the bottom half, often with some median wall thickening, the spores with a majority that are weakly ornamented (O2), albeit some may be more strongly ornamented or almost smooth and the perispore which is not consistently or distinctly loosening (rarely P2 and never P3) and the pileus colour which always has brown or buff tones at least in the centre. Hebeloma hiemale most likely corresponds to ICG21 in the intercompatibility tests of Aanen & Kuyper (1999). In Beker et al. (2010) we discussed the difficulty in interpreting the precise taxon to which this name should refer. Hebeloma hiemale was described by Bresadola in 1898 and the lectotype is an icon from Fung. Trident. Pl.160. It is likely that Bresadola’s concept of this species embraced more than one taxon. As we described in Beker et al. (2010), there are two collections labelled H. hiemale in the Swedish Museum of Natural History, both collected by Bresadola in Gocciadoro, Trentino, which is the location cited in the original 1898 description. Collection F14406 was collected in November 1914. A microscopic analysis reveals spores and cheilocystidia typical of H. subsect. Crustuliniformia and the presence of a large number of full length lamellae would indicate this as being one of the larger species within this group. Further, the spores are quite short, averaging 10.6 μm, and so this collection would be highly unlikely to resemble Bresadola’s icon or original description. The average cheilocystidium apex width for this collection is 7.3 μm and so this collection can be referred to H. aanenii. Collection F28516 was collected in November 1900. This collection is mixed, as noted by E. Grilli (herbarium note). We hypothesised (Beker et al. 2010) that part of the collection likely represents H. cavipes, (H. subsect. Clepsydroida) a species similar to H. hiemale but with more ornamented and more dextrinoid spores, usually on average larger than those of H. hiemale. The spores of this collection have an average length of 13.2 μm and a Q greater than 2. We have since generated molecular evidence (ITS sequence) that this part of the collection can indeed be assigned to H. cavipes. The second part of the collection is more difficult to determine. Although many of the cheilocystidia are very badly collapsed, we were able to find some in the fragment we examined, and this, together with the spores (ave. 11.1 × 5.9 μm), does indicate that this part of the collection should be assigned to H. hiemale. Our database query for H. hiemale does capture this part of the collection as being H. hiemale. To resolve the ambiguity around the precise definition of H. hiemale and to provide nomenclatural stability, we selected an epitype for H. hiemale which ties it to a single taxon. In Beker et al. (2010), we discussed the idea that if it were possible to sequence one of Bresadola’s collections that had been studied by Grilli (1997), this could be an ideal collection for epitypification. Unfortunately (Grilli, private comm.), to date no sequence (ITS or otherwise) has been forthcoming from this material. As a consequence we have selected a different collection as epitype, that in our view exhibits all the properties of Hebeloma hiemale (Eberhardt et al. 2016).

Geographic distribution

Phenology

• arrow_drop_downarrow_drop_upAdditional cited collections

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