Hebeloma velutipes (Photo: J. Vesterholt)

Taxonomy

• arrow_drop_downarrow_drop_upNomenclatural notes
  It has been proposed that the name Hebeloma velutipes Bruchet be conserved against H. bakeri Earle by Beker et al., (2793) Proposal to conserve the name Hebeloma velutipes against H. bakeri (Fungi, Agaricales, Hymenogastraceae) in Taxon 70: 202-203 (2021),

Types: FRANCE: Les Echets (Ain) (approx. 45.86°N, 4.9°E, alt. approx. 280 m a.s.l.) on clayey soil in deciduous woodland under Corylus avellana, 24 Sep. 1963, G. Bruchet (Holotype. herbarium acc. no. LY BR63-20, HJB1000045).

Heterotypic synonyms:

• Hebeloma angustifolium Romagn., Sydowia 36: 258 (1947) ["1983"]
• Hebeloma bulbosum Romagn., Sydowia 36: 263 (1947) ["1983"]
• Hebeloma favrei Romagn. & Quadr., Documents mycologiques 14 (56): 31 (1985) ["1984"]
• Hebeloma mediterraneum (A. Gennari) Contu, Bollettino della Associazione Micologica ed Ecologica Romana 73-74 (1-2): 16 (2008)
• Hebeloma stenocystis J. Favre ex Quadr., Mycol. Helv. 3 (2): 197 (1989) ["1988"]
• Hebeloma tenuifolium Romagn., Documents mycologiques 15 (59): 53 (1985)
• Hebeloma bakeri Earle nom. rej. prop., Bulletin of the New York Botanical Garden 2: 342 (1902)
• Hebeloma latisporum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 142 (1983)
• Hebeloma pallidoargillaceum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 146 (1983)
• Hebeloma salmonense A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 86 (1983)
• Hebelomina mediterranea A. Gennari, Rivista di Micologia 4: 312 (2002)

Homotypic synonyms:

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

• arrow_drop_downarrow_drop_upEtymology
  From the adjective velutinus– velvety and the noun pes– foot to emphasise the stipitipellis as being velutinous, i.e. the velvet-like appearance of the stipe surface due to the dense covering of caulocystidia.

• arrow_drop_downarrow_drop_upOriginal diagnosis
  Cortina nulla; pileo majore, usque ad 50 mm lato, convexo vel e plano convexo, haud mammoso, concolore, jove sicco, luteolo, ex ochraceo luteo vel e gilvo luteo, parum carnoso; stipite 50–70 x 6–8 mm, longiore, haud gracili, subbulboso, albido, sub lamellis velutino ac inferius passim floccoso; lamellis guttulis lacteis in acie madidis; odore debiliter raphanoideo. Sporis 10–12 x 5,5–6 μm, amygdaliformibus, brevioribus, parum ventricosis, s.m. brunneis, verrucosis, pariete firma; pilis marginum 50–65 μm longis, summis ad 7,5–10 μm capitatis. Sub Betulis Salicibus mixtis.

• arrow_drop_downarrow_drop_upEnglish translation
  Cortina absent; pileus rather large, up to 50 mm broad, convex or plano-convex, non-umbonate, of a uniform colour, in dry weather ochraceous-yellow to pale yellowish or pale yellow to clay, hardly fleshy; stipe 50–70 × 6–8 mm, rather long, not at all gracile, sub-bulbous, whitish, velutinous below lamellae and here and there floccose lower down; lamella edge exuding milky beads; odour faintly raphanoid. Spores 10–12 × 5.5–6 μm, amygdaloid, rather short, hardly ventricose, under the microscope brown, verrucose, thick-walled; marginal hairs 50–65 μm long, with a 7.5–10 μm wide capitate apical part. Under mixed Betula and Salix.

Description

Description of Hebeloma velutipes based on 892 collections

• arrow_drop_downarrow_drop_upMacroscopic description
  Pileus: (12) 20–64 (95) mm diameter; shape occasionally convex, umbonate or broadly umbonate, rarely strongly umbonate, weakly umbonate, applanate or umbilicate; characters occasionally spotting, rarely hygrophanous, remains of universal veil, rugulose or rimulose; margin characters usually smooth, rarely involute, reflexed, overhanging pileus, wavy, fibrillose, ribbed or scalloped; viscosity tacky when moist; colour variation often unicolour, occasionally two color; colour at centre occasionally cream, rarely warm buff, pale cream, pale yellow, pinkish buff, dark pinkish buff, honey, pale pinkish buff, buff-yellow, ochraceous, yellowish brown, cinnamon, umber, clay-buff, brownish olive or clay-red.

  Lamellae: attachment usually emarginate, rarely adnate, adnexed, decurrent tooth, decurrent or free; maximum depth 2–64 mm; number of complete lamellae 41–80; presence of tears often visible with naked eye, occasionally absent or visible with x10 lens; white fimbriate edge usually present, rarely very strong, weak or absent.

  Cortina presence: usually no, rarely yes.

  Stipe: (5) 29–80 (120) x (3) 4–10 (20) {median} x (3) 5–16 (27) {basal} mm; stipe Q 0.6–22.5; base shape often clavate, occasionally bulbous or cylindrical, rarely subbulbous or tapering; floccosity occasionally floccose, pruinose or velute, rarely pruinose at apex, weakly floccose, fibrillose, floccose at apex, fibrous, none or powdery; rooting no; thick rhizoids at base usually absent, rarely present;

  Context: Texture firm; stipe interior usually hollow, occasionally superior wick or stuffed, rarely basal wick; stipe flesh discolouring often no, occasionally yes, rarely weak or very strongly; slenderness measure 1.5–41.9; smell often raphanoid, rarely strongly raphanoid, weakly raphanoid, cocoa, odourless, earthy, fruit, soap or sweet; taste occasionally raphanoid, bitter or mild, rarely hot, weakly bitter, weakly raphanoid, strongly bitter or strongly raphanoid where recorded.

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

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

• arrow_drop_downarrow_drop_upMicroscopic description
  Spores: shape usually amygdaloid, occasionally limoniform, rarely lacrymoid; colour in microscope often yellow brown, occasionally yellow or brown, rarely brown yellow, grey yellow or very pale; guttules often yes, occasionally no, rarely weak. papilla variable occasionally weak, rarely very strongly; Spore Code: O2 (O3); P0 P1; (D2) D3.

  Basidia: (21) 24–37 (42) x (5) 6–9 (10) μm; ave. Q 3.0–4.9; spore arrangement 4 spored;

  Cheilocystidia: main shape gently clavate, often clavate-lageniform or clavate-ventricose, occasionally ventricose, clavate-stipitate or clavate, rarely cylindrical, lageniform, tapering, capitate-stipitate, capitate, subcapitate or utriform; special features observed occasionally bifurcate or septa, rarely geniculate, apical thickening, clamped septa, branching, median thickening, plaques, sinuate, mucous, conglutinate, irregular, many collapsed in exsiccata, short, sparse, tapering or yellow contents; cheilocystidia ratios: A/M = 1.19–1.83; A/B = 1.07–2.28; B/M = 0.73–1.34.

  Pleurocystidia: none seen.

  Ixocutis: epicutis thickness (measured from exsiccata) up to 200 μm; ixocutis hyphae width up to 8 μm; ixocutis hyphae encrustation usually yes, rarely no; shape of trama elements beneath subcutis occasionally cylindrical or thickly sausage-shaped, rarely ellipsoid, oblong, polygonal, thinly sausage-shaped, angular, hyphae thin, isodiametric or spherical up to 16 μm wide.

  Caulocystidia: Similar to cheilocystidia but larger, up to 200 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 Betula (14.2%) but Pinus (12.9%), Picea (11.9%), Quercus (10.6%), Salix (9.9%), Fagus (8.0%), Dryas (4.3%), Abies (4.3%), Tsuga (4.0%), Pseudotsuga (3.6%), Helianthemum (2.6%), Carpinus (2.6%), Populus (2.0%), Corylus (2.0%), Cedrus (1.3%), Tilia (1.3%) and Notholithocarpus (1.3%) were also recorded. In these cases the most commonly recorded families were Pinaceae (48.5%), Fagaceae (18.0%), Betulaceae (15.7%) and Salicaceae (10.3%). We have additional records where Alnus (4.7%), Arbutus (2.0%) and Larix (1.6%) were recorded as possible associates, but in these cases a number of possible associates were mentioned. Overall the most commonly recorded families are Pinaceae (56.4%), Betulaceae (33.1%), Fagaceae (27.3%) and Salicaceae (25.8%) The growth habit of our collections was often scattered, occasionally solitary and rarely gregarious, caespitose or connate.

  According to our current data, the species is found on multiple continents with collections found in Europe (50.5%), Northern America (40.8%), Temperate Asia (5.6%), Australasia (3.0%) and Southern America (0.1%). On these continents, collections have 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 (37.9%), temperate conifer forests (27.8%) and boreal forests/taiga (12.7%). From collector information, it appears collections have been found in the 1.4 Forest – Temperate (57.1%) and 1.1 Forest – Boreal (12.7%) 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 Europe we have records from the Centre (Belgium, Germany, Poland, Slovakia, Switzerland, Netherlands and Czech Republic), the North (England, Norway, Svalbard, Scotland, Denmark, Isle Of Man, Iceland, Sweden, Wales, Finland, Faroe Islands and Ireland), the Southwest (France, Spain, Italy, Portugal and Andorra), the Southeast (Italy, North Macedonia and Bulgaria) and Eastern Europe (Lithuania). Specimens have been collected from 32.8°N to 78.9°N.

  Within Northern America we have records from Subarctic America (Alaska, Northwest Territories, Yukon and Greenland), Northwestern U.S.A. (Washington, Colorado, Oregon, Montana and Idaho), Southwestern U.S.A. (California and Arizona), Western Canada (British Columbia, Alberta and Manitoba), Eastern Canada (Quebec, Newfoundland and Labrador, New Brunswick, Nova Scotia and Ontario), Mexico (Mexico) and Northeastern U.S.A. (Massachusetts, New Jersey and Michigan).

  Within Temperate Asia we have records from Western Asia (Cyprus and Turkey), Caucasus (Karachay-Balkar, Georgia and Adygea), Russian Far East (Kamchatka, Chukotka and Magadanskaya), Eastern Asia (Japan), Siberia (Tyumen and Krasnoyarsk) and China (Sichuan).

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

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

• arrow_drop_downarrow_drop_upMolecular results
  Aanen et al. (2001) showed that the intragenomic ITS variation found in some collections (genomes) of H. velutipes is of an unusually high magnitude. These collections contain two different ITS copies that are so different that they are not even contained in sister groups in cladistic analyses. These clades correspond to H. incarnatulum on one hand and H. leucosarx and H. subconcolor on the other (Grilli et al. 2016). Aanen and co-workers (2001) also showed that these ITS copies can be segregated in monokarya. Molecular recognition of Hebeloma velutipes is difficult and none of the loci tested can on its own be used to unambiguously identify this taxon. The concatenation of all loci is possibly less powerful then analysing the results of the different loci separately. A combination of several loci could then be used to exclude all other taxa from the velutipes-complex. Habitat or geographical information could also be used to narrow down further the number of possible taxa in the identification process. The ITS situation suggests that the evolutionary history of H. velutipes was influenced by reticulate processes and, thus, it cannot not properly be reconstructed by tree-building methods. The synonymization of several species, H. favrei, H. mediterraneum, H. stenocystis and H. tenuifolium is supported by ITS and (apart from H. favrei) also by V6 and V9 sequence data (Grilli et al. 2016).

• arrow_drop_downarrow_drop_upCommentary
  With its gently clavate cheilocystidia, interspersed with some cheilocystidia that are clavate-lageniform or ventricose and the spores rather strongly dextrinoid, this taxon clearly belongs to H. sect. Velutipes. Within this section, the cheilocystidium ratio B/M being less than 1.35 and the spores rarely a combination of distinctly ornamented, perispore distinctly and strongly loosening and strongly dextrinoid (i.e. not O3, P2.P3, D4) takes us to members of the velutipes-complex, H. leucosarx, H. incarnatulum and H. velutipes. Microscopically, the cheilocystidia of H. incarnatulum are far more slender and the average apex of the cheilocystidia never exceeds 6.5 μm. The average apex of the cheilocystidia in H. velutipes ranges from 6.4–8.9 μm and is usually well over 7 μm. Indeed, with over 700 collections of this taxon, we have only twice recorded an average apex value of 6.4 μm and twice an average value of 6.5 μm. Microscopically, we have not found any way to separate H. leucosarx from H. velutipes, however, macroscopically, they are quite different. Hebeloma leucosarx is always distinctly umbonate, with a slender appearance and a dark coloured pileus. Hebeloma velutipes has a much paler pileus colour (it is very rarely dark) and usually a less slender appearance; it too can be quite distinctly umbonate. Also, the exsiccata of these two taxa is quite different: H. leucosarx is always a rich brown colour while H. velutipes has a distinctive pallid matt look to the pileus of dried material. While it is difficult to describe the characteristics of the exsiccata, it is very distinctive and usually readily recognizable. Phylogenetically, H. velutipes appears to be closest to H. incarnatulum and H. subconcolor, but these taxa can usually be easily separated macroscopically and on the basis of habitat (and are microscopically quite different as discussed above). Hebeloma incarnatulum is invariably found in Sphagnum with conifers while H. velutipes is rarely in such a habitat. Of the 304 collections on which we based our habitat and distribution discussion, only two records mentioned the presence of Sphagnum and, as it happens, neither of these two recorded conifers. Hebeloma subconcolor is a much smaller fungus than H. velutipes and with far fewer lamellae. Not surprisingly, H. velutipes has a number of synonyms (and probably many more than we list above, but in many cases we cannot rule out some ambiguity and hence remain cautious). These synonyms include Hebeloma mediterraneum, which was first described as Hebelomina mediterranea (the spores of this collection were omitted from our composite species description.) Hebeloma velutipes can be very variable, both from a molecular perspective and morphologically. For example, the spores are very variable in size, ornamentation, dextrinoidity and so on. Currently we ‘lump’ all these together in the taxon we call H. velutipes, but it is possible that this represents a complex of species. While we have looked for differences that are consistent and with some support molecularly, so far we have not found sufficient evidence for separation. However, as mentioned above, we do have collections from outside Europe that we do know represent other taxa close to H. velutipes and within H. sect. Velutipes. Hebeloma velutipes is, alongside H. cavipes and H. mesophaeum, one of the most common ebeloma sp. in Europe. At the time of writing, collections of H. velutipes represent over 9% of the 4500 collections on our database.

Geographic distribution

Phenology

• arrow_drop_downarrow_drop_upAdditional cited collections

Image gallery