Hebeloma alpinicola (Photo: E. Campo)

Taxonomy

Types: UNITED STATES: Idaho: Heavens Gate Ridge, Seven Devils Mountains (approx. 45.3688°N, 116.4938°W, alt. approx. 2560 m a.s.l.) in coniferous, subalpine woodland under Pinus albicaulis, 5 Jul. 1958, A.H. Smith (58632) (Holotype. herbarium acc. no. MICH 5549, HJB1000311).

Heterotypic synonyms:

• Hebeloma angustifolium A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 164 (1983)
• Hebeloma brunneodiscum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 39 (1983)
• Hebeloma chapmaniae A.H. Sm., V.S. Evenson & Mitchel [as "chapmanae"], The Veiled Species of Hebeloma in the Western United States: 37 (1983)
• Hebeloma dissiliens A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 60 (1983)
• Hebeloma glabrescens A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 65 (1983)
• Hebeloma griseocanescens A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 50 (1983)
• Hebeloma littenii A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 169 (1983)
• Hebeloma mesophaeum var. aspenicola A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 76 (1983)
• Hebeloma mesophaeum var. castaneum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 77 (1983)
• Hebeloma mesophaeum var. duplicatum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 79 (1983)
• Hebeloma mesophaeum var. fluviatile A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 81 (1983)
• Hebeloma mesophaeum var. insipidum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 80 (1983)
• Hebeloma mesophaeum var. similissimum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 83 (1983)
• Hebeloma nigromaculatum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 51 (1983)
• Hebeloma perigoense A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 57 (1983)
• Hebeloma pseudofastabile var. distans A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 114 (1983)
• Hebeloma riparium A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 43 (1983)
• Hebeloma sanjuanense A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 62 (1983)
• Hebeloma smithii Quadr., Mycotaxon 30: 303 (1987)
• Hebeloma solheimii A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 68 (1983)
• Hebeloma strophosum var. occidentale A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 63 (1983)
• Hebeloma subannulatum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 42 (1983)
• Hebeloma subargillaceum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 58 (1983)
• Hebeloma subcapitatum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 67 (1983)
• Hebeloma sublamellatum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 55 (1983)
• Hebeloma subrimosum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 26 (1983)
• Hebeloma substrophosum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 45 (1983)
• Hebeloma subviolaceum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 66 (1983)
• Hebeloma vinaceoumbrinum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 46 (1983)

• arrow_drop_downarrow_drop_upEtymology
  a dweller of alpine areas.

• arrow_drop_downarrow_drop_upOriginal diagnosis
  Pileus 2–4 cm latus, demum subplanus vel undulatus, ad marginem fibrillosus, glabrescens, glutinosus, cinnamomeus. Odor et gustus raphaninus. Lamellae subdistantes, latae, demum fulvocinnamomeae. Stipes 3–6 cm longus, 4–8 (11) mm crassus, aequalis, albidus tarde brunnescens. Velum bicoloratum (ochraceum et albidum). Sporae 7–9 x 5–5.5 μm, ovoideae vel ellipsoideae, non dextrinoideae. Cheilocystidia 28–43 x 5–7μm, fusoid-ventricosa vel clavata vel subcylindrica. Specimen typicum in Herb. Univ. Mich. conservatum est, Smith 58632; legit sub Pinus albicaulis, prope Riggins, Idaho, 5 Jul 1958.

• arrow_drop_downarrow_drop_upEnglish translation
  Pileus 2–4 cm broad, finally subapplanate or undulate, fibrillose at margin, becoming glabrous, cinnamon, yellowish-brown.. Odour and taste raphanoid. Lamellae subdistant, broad, finally cinnamon-fulvous. Stipe 3–6 cm long, 4–8 (11) mm thick, equal, whitish becoming slowly brown. Veil two-coloured (ochraceous and whitish). Spores 7–9 x 5–5.5 μm, ovoid or ellipsoid, indextrinoid. Cheilocystidia 28–43 x 5–7 μm, fusoid-ventricose, clavate or cylindrical.

Description

Description of Hebeloma alpinicola based on 158 collections

• arrow_drop_downarrow_drop_upMacroscopic description
  Pileus: (7) 17–44 (70) mm diameter; shape often convex, occasionally umbonate or broadly umbonate, rarely strongly umbonate, umbilicate, weakly umbonate or none; characters often remains of universal veil, rarely spotting, hygrophanous or pubescent; margin characters often involute, rarely fibrillose, smooth, reflexed or wavy; viscosity tacky when moist; colour variation often unicolour, occasionally two color; colour at centre occasionally cinnamon, yellowish brown or brownish olive, rarely fuscous, clay-buff, umber, sepia, dark pinkish buff, orange-brown or ochraceous.

  Lamellae: attachment often adnate, occasionally emarginate, rarely adnexed, decurrent tooth or free; maximum depth 2–5 mm; number of complete lamellae 30–48; presence of tears usually absent, rarely visible with x10 lens; white fimbriate edge occasionally present, weak or absent.

  Cortina presence: yes.

  Stipe: (15) 22–73 (100) x (1) 2–10 (13) {median} x (1) 3–10 (13) {basal} mm; stipe Q 2.9–25.0; base shape usually cylindrical, rarely clavate or tapering; floccosity usually fibrillose, rarely pruinose at apex, fibrous, floccose at apex or velute; rooting no; thick rhizoids at base absent;

  Context: Texture firm; stipe interior often hollow, occasionally stuffed, rarely superior wick; stipe flesh discolouring often yes, rarely no, very strongly or weak; slenderness measure 3.0–51.3; smell occasionally raphanoid or odourless, rarely weakly raphanoid, cocoa or strongly raphanoid; taste often raphanoid, occasionally mild, rarely none, weakly bitter or bitter where recorded.

  Spore deposit colour: often clay-buff, occasionally sepia or yellowish brown.

  Exsiccata characters: occasionally dark.

• arrow_drop_downarrow_drop_upMicroscopic description
  Spores: shape usually ellipsoid, often ovoid, rarely cylindrical or amygdaloid; colour in microscope often yellow or yellow brown, rarely very pale, yellow pale, brown or grey yellow; guttules often no, occasionally yes, rarely weak. papilla no; Spore Code: O1; P0; D0 (D1).

  Basidia: (18) 19–33 (36) x (5) 6–8 (9) μm; ave. Q 2.9–4.5; spore arrangement 4 spored;

  Cheilocystidia: main shape ventricose, usually lageniform, occasionally cylindrical, rarely clavate-lageniform or clavate-ventricose, balloon-shaped, capitate, clavate, gently clavate, lanceolate or pyriform; special features observed often septa, occasionally many collapsed in exsiccata or geniculate, rarely yellow contents, basal thickening, median thickening, short, subcapitate, thick content in neck, bifurcate, branching, clamped septa, grotesque, irregular or rostrate; cheilocystidia ratios: A/M = 0.95–1.44; A/B = 0.54–0.90; B/M = 1.43–2.19.

  Pleurocystidia: usually none seen, rarely seen or only close to lamella edge.

  Ixocutis: epicutis thickness (measured from exsiccata) up to 200 μm; ixocutis hyphae width up to 8 μm; ixocutis hyphae encrustation no; shape of trama elements beneath subcutis cylindrical.

  Caulocystidia: Similar to cheilocystidia but larger.

• arrow_drop_downarrow_drop_upSpore measurements

• arrow_drop_downarrow_drop_upCheilocystidia measurements

• arrow_drop_downarrow_drop_upHabitat and distribution
  Hebeloma alpinicola's preferred substrate appears to be acidic soil. Where only one possible associate was recorded, the most commonly recorded associate was Salix (37.3%) but Picea (28.8%), Pinus (11.9%), Dryas (6.8%), Betula (5.1%), Populus (5.1%), Alnus (3.4%) and Pseudotsuga (1.7%) were also recorded. In these cases the most commonly recorded families were Pinaceae (50.8%), Salicaceae (34.4%), Betulaceae (8.2%) and Rosaceae (6.6%). We have additional records where Arctostaphylos (3.5%), Abies (2.6%) and Tsuga (2.6%) were recorded as possible associates, but in these cases a number of possible associates were mentioned. Overall the most commonly recorded families are Salicaceae (46.5%), Pinaceae (37.7%), Betulaceae (29.8%) and Rosaceae (7.9%) The growth habit of our collections was often scattered, occasionally gregarious and rarely solitary or caespitose.

  According to our current data, the species is found on multiple continents with collections found in Northern America (67.5%), Europe (20.1%) and Temperate Asia (12.3%). 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 conifer forests (50.0%), tundra (17.1%), temperate broadleaf & mixed forests (16.5%) and boreal forests/taiga (10.8%), specifically including the ecoregions: Colorado Rockies forests (24.7%) and Caucasus mixed forests (11.4%). From collector information, it appears collections have been found in the 1.1 Forest – Boreal (40.1%), 3.3 Shrubland – Boreal (17.5%), 5.11 Wetlands (inland) – Alpine wetlands (inc. temporary waters from snowmelt) (16.8%) and 4.1 Grassland – Tundra (11.7%) 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 Northwestern U.S.A. (Colorado, Washington, Idaho, Montana and Oregon), Subarctic America (Alaska, Greenland, Northwest Territories and Yukon), Southwestern U.S.A. (California and Arizona), Eastern Canada (Newfoundland and Labrador and Quebec), Northeastern U.S.A. (Michigan and Maine), North-central U.S.A. (Minnesota), Western Canada (British Columbia) and South-central U.S.A. (New Mexico).

  Within Europe we have records from the North (Norway, Iceland, Faroe Islands, Sweden, Finland and Scotland), the Southeast (Italy and Romania), the Southwest (France) and the Centre (Austria and Switzerland). Specimens have been collected from 42.7°N to 69.9°N.

  Within Temperate Asia we have records from Caucasus (Karachay-Balkar and Georgia) and Russian Far East (Kamchatka).

• arrow_drop_downarrow_drop_upCommentary
  This taxon, with its small ellipsoid, indextrinoid spores and ventricose cheilocystidia is a member of H. sect. Hebeloma. Morphologically it is closely related to H. excedens and H. mesophaeum. It is generally more robust than these two species, especially the stipe, and the pileus is not as two-toned. While further work is needed to decide whether this really is a species distinct from the other two, the molecular evidence coupled with the morphological evidence suggest this to be the case. We have studied a number of collections, from a variety of habitats within North America that all appear to represent this taxon. A number of species representing this taxon were published by Smith et al. (1983) in the same publication that featured H. alpinicola. Although there is some molecular variation between these different collections, it is very small and we see insufficient evidence to separate these species. We have selected the name Hebeloma alpinicola on the grounds that although not all collections are strictly alpine, the majority are at least subalpine.

Geographic distribution

Phenology

• arrow_drop_downarrow_drop_upAdditional cited collections

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