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Taxonomy

Full name: Hebeloma palustre Peck, Ann. Rep. N.Y. St. Mus. 52: 649 (1899)
Genus: Hebeloma
Section: Hebeloma
Subsection: 'subsect1'

Types: UNITED STATES: New York: Kasoag (approx. 43.46°N, 75.92°W, alt. approx. 190 m a.s.l.) on boggy, mossy soil in bog, Oct. 1898, C.H. Peck (Lectotype. herbarium acc. no. NYS-F-002244.1, HJB1000088). Lectotype designated by Eberhardt et al., Mycologia 114 (2): (2022) page 362 (MBT10000878).

Heterotypic synonyms:
  • Hebeloma clavulipes Romagn., Bull. Trimestriel Soc. Mycol. France 81 (3): 326 (1929) ["1965"]
  • Hebeloma oreophilum Beker & U. Eberh., Mycologia 107 (6): 1295 (2016) ["2015"]
  • Hebeloma candidipes Bruchet, Bull. Mens. Soc. Linn. Lyon 39, supplement 6: 125 (1970)
  • Hebeloma mesophaeum var. macrosporum Remy, Bulletin trimestriel de la Société Mycologique de France 80 (4): 531 (1965) ["1964"]
  • Hebeloma remyi Bruchet, Bull. Mens. Soc. Linn. Lyon 39, supplement 6: 29 (1970)
  • Hebeloma corrugatum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 116 (1983)
  • Hebeloma discomorbidum (Peck) Peck, Bull. N.Y. St. Mus.: 75 (1910)
  • Agaricus discomorbidus Peck [as "Agaricus (Naucoria) discomorbidus"], Bull. Buffalo Soc. nat. Sci. 1: 52 (1873)
  • Hebeloma felleum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 151 (1983)
  • Hebeloma flaccidum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 144 (1983)
  • Hebeloma fragrans A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 91 (1983)
  • Hebeloma fragrans var. intermedium A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 92 (1983)
  • Hebeloma fuscostipes A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 119 (1983)
  • Hebeloma griseocanum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 113 (1983)
  • Hebeloma hesleri A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 122 (1983)
  • Hebeloma idahoense A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 143 (1983)
  • Hebeloma limacinum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 103 (1983)
  • Hebeloma lutescentipes A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 154 (1983)
  • Hebeloma marginatulum var. fallax A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 130 (1983)
  • Hebeloma obscurum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 124 (1983)
  • Hebeloma occidentale A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 121 (1983)
  • Hebeloma oregonense var. atrobrunneum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 150 (1983)
  • Hebeloma pallescens A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 137 (1983)
  • Hebeloma perfarinaceum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 153 (1983)
  • Hebeloma piceicola A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 126 (1983)
  • Hebeloma praecaespitosum A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 118 (1983)
  • Hebeloma praelatifolium A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 157 (1983)
  • Hebeloma pseudofastabile A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 127 (1983)
  • Hebeloma stanleyense A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 147 (1983)
  • Hebeloma subboreale A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 33 (1983)
  • Hebeloma subrubescens A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 136 (1983)
  • Hebeloma utahense A.H. Sm., V.S. Evenson & Mitchel, The Veiled Species of Hebeloma in the Western United States: 29 (1983)
  • Hygrophorus montanus Murrill, Mycologia 3 (4): 199 (1911)

  • arrow_drop_downarrow_drop_upEtymology
    From palustris, meaning ‘swampy, marshy’.
  • arrow_drop_downarrow_drop_upDiagnosis
    Pileus fleshy but thin, broadly convex becoming nearly plane with age, sometimes wavy or irregular, glabrous, hygrophanous, grayish brown and slightly striatulate on the margin when moist, paler when dry, flesh whitish; lamellae close thin ventricose, adnexed, grayish white becoming cinnamon brown; stem, rather long, equal or tapering upward, hollow, silky, white; spores subelliptic, uninucleate, .0004 to .0005 in. long, .00024 to .0003 broad [10.2-12.7 x 6.1-7.6 µm]. Pileus 1 to 1.5 in. broad [25.4-38.1 mm]; stem 2 to 3 in. long [50.1-76.2 mm], 2 to 4 lines thick [5.1-10.2 mm]. Mossy ground in swampy woods. Kasoag. October. The pileus is not viscid and there is no evidence of a veil.

Description

  • arrow_drop_downarrow_drop_upThresholds
Description of Hebeloma palustre based on 140 collections
  • arrow_drop_downarrow_drop_upMacroscopic description
    Pileus: (8) 15–35 (50) mm diameter; shape usually convex, occasionally umbonate, rarely weakly umbonate, strongly umbonate, applanate or broadly umbonate; characters often remains of universal veil, occasionally hygrophanous; margin characters often smooth, occasionally involute or fibrillose, rarely sulcate, reflexed, wavy or crenulate; viscosity tacky when moist; colour variation often two color, occasionally unicolour; colour at centre occasionally brownish olive, rarely sepia, umber, yellowish brown, fuscous, cinnamon, dark fawn, dark brick, brick, clay-buff, orange-brown, fawn, greyish brown, ochraceous or cream.

    Lamellae: attachment often emarginate, occasionally adnate, rarely adnexed; maximum depth 2–8 mm; number of complete lamellae 38–50; presence of tears usually absent, rarely visible with x10 lens or visible with naked eye; white fimbriate edge often present, occasionally weak, rarely absent.

    Cortina presence: yes.

    Stipe: (13) 24–69 (110) x (1) 2–6 (10) {median} x (1) 2–7 (12) {basal} mm; stipe Q 3.3–25.0; base shape usually cylindrical, occasionally clavate, rarely tapering or bulbous; floccosity often fibrillose, occasionally pruinose at apex, rarely floccose at apex, pruinose, fibrous, floccose or none; rooting no; thick rhizoids at base absent;

    Context: Texture firm; stipe interior often hollow, occasionally stuffed; stipe flesh discolouring often yes, rarely no, weak or very strongly; slenderness measure 4.2–645.3; smell often raphanoid, rarely strongly raphanoid, odourless, weakly raphanoid, earthy, cocoa or fruit; taste occasionally mild or raphanoid, rarely weakly bitter, none, strongly bitter, weakly raphanoid, bitter or strongly raphanoid where recorded.

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

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

  • arrow_drop_downarrow_drop_upMicroscopic description
    Spores: shape amygdaloid, occasionally limoniform, rarely ovoid or fusoid; colour in microscope occasionally yellow brown, yellow or grey yellow, rarely brown, brown pale or brown yellow; guttules often no, occasionally yes, rarely weak. papilla often yes, occasionally weak, rarely no or very strongly; Spore Code: (O1) O2; P0 (P1); D2 D3.

    Basidia: (17) 21–36 (38) x (5) 6–10 μm; ave. Q 2.8–4.3; spore arrangement 4 spored;

    Cheilocystidia: main shape usually lageniform or ventricose, occasionally cylindrical, rarely pyriform, balloon-shaped, clavate or clavate-lageniform or clavate-ventricose; special features observed often septa, occasionally median thickening, basal thickening, geniculate or many collapsed in exsiccata, rarely apical thickening, clamped septa, sparse, thick content in neck or yellow contents; cheilocystidia ratios: A/M = 0.87–1.46; A/B = 0.42–0.81; B/M = 1.49–2.40.

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

    Ixocutis: epicutis thickness (measured from exsiccata) up to 170 μm; ixocutis hyphae width up to 7 μm; ixocutis hyphae encrustation often no, occasionally yes; shape of trama elements beneath subcutis often thickly sausage-shaped, occasionally thinly sausage-shaped, rarely cylindrical or polygonal up to 18 μm wide.

    Caulocystidia: Similar to cheilocystidia but larger, up to 120 μm.

  • arrow_drop_downarrow_drop_upSpore measurements
  • arrow_drop_downarrow_drop_upCheilocystidia measurements
  • arrow_drop_downarrow_drop_upHabitat and distribution
    Hebeloma palustre's preferred substrate appears to be mossy soil. Where only one possible associate was recorded, the most commonly recorded associate was Salix (60.9%) but Picea (25.0%), Betula (3.1%), Alnus (3.1%), Fagus (1.6%), Dryas (1.6%), Populus (1.6%), Tsuga (1.6%) and Pinus (1.6%) were also recorded. In these cases the most commonly recorded families were Salicaceae (51.4%), Pinaceae (41.7%) and Betulaceae (5.6%). We have additional records where Abies was recorded as a possible associate, but for these collections a number of possible associates were mentioned. Overall the most commonly recorded families are Salicaceae (57.7%), Pinaceae (38.7%) and Betulaceae (24.3%) The growth habit of our collections was often scattered, occasionally gregarious and rarely caespitose or solitary.

    According to our current data, the species is found on multiple continents with collections found in Northern America (55.4%), Europe (40.3%), Temperate Asia (3.6%) and Southern America (0.7%). 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 (44.8%), tundra (24.6%) and temperate broadleaf & mixed forests (11.2%), specifically including the ecoregions: Colorado Rockies forests (23.1%) and Kalaallit Nunaat Arctic steppe (11.2%). From collector information, it appears collections have been found in the 1.1 Forest – Boreal (31.1%), 4.1 Grassland – Tundra (25.4%), 1.4 Forest – Temperate (12.3%) and 5.11 Wetlands (inland) – Alpine wetlands (inc. temporary waters from snowmelt) (11.5%) 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 Northern America we have records from Northwestern U.S.A. (Colorado, Montana, Idaho, Oregon, Wyoming and Washington), Subarctic America (Greenland, Alaska and Northwest Territories), Northeastern U.S.A. (New York), Eastern Canada (Newfoundland and Labrador and Quebec) and Southwestern U.S.A. (Utah).

    Within Europe we have records from the North (Svalbard, Finland, Sweden, Norway, Denmark and England), the Centre (Slovakia, Germany, Poland and Czech Republic), the Southwest (France) and the Southeast (Italy). Specimens have been collected from 44.9°N to 78.3°N.

    Within Temperate Asia we have records from Siberia (Krasnoyarsk and Tyumen) and Russian Far East (Sakha).

    Within Southern America all our records are from Caribbean (Jamaica).

  • arrow_drop_downarrow_drop_upCommentary
    Hebeloma palustre belongs to H. sect. Hebeloma given the ventricose to lageniform cheilocystidia together with the amygdaloid and limoniform spores. Given the habitat, it is most likely in the group of species that includes H. clavulipes, H. hygrophilum, H. monticola and H. nigellum. The protologue mentions that the lamellae are close and from the exsiccata it would appear to be between 44 and 48, although estimating from the exsiccata is never without risk; this would rule out H. hygrophilum and H. nigellum, which normally never have more than 36 full-length lamellae. The spores, while a little short for H. clavulipes, could correspond to either H. clavulipes or H. monticola, but the number of lamellae would fit better with H. clavulipes. While we were unable to generate any sequences from the material, that might confirm this identification, we are confident that this is a correct identification. Note that H. discomorbidum, another species described by Peck, is conspecific. The name H. palustre has priority over both H. clavulipes and H. discomorbidum as well as over numerous other names, the majority of which were published by Smith et al. (1983). Twenty of the types of taxa published by Smith, synonymized with H. palustre above, were collected in Pitkin county, Colorado, between the years (1976) 1978–1980. It should be noted that we did not receive any type material for H. fragrans var. intermedium because of the sparsity of the material. Instead, we examined a topotype collected on the same day. Among the many heterotypic synonyms of H. palustre we now also include H. oreophilum. This latter taxon was originally described from the Western Carpathian Mountains in Slovakia. In Beker et al. (2016) it was pointed out that this species was both morphologically and molecularly closely related to H. clavulipes but could be separated on both habitat and its wider spores. Its description was based on 13 collections, while that of H. clavulipes was based on 21 collections. Since that publication we have had the opportunity to study many more collections of both taxa, including many collections from North America. We must conclude that this is a single species with a continuum of average spore width from 6–7.5 µm, and a range of habitats from lowland to arctic/alpine. This is similar to species such as Hebeloma dunense (but see H. velatum below), H. mesophaeum and H. nigellum which, as discussed in Beker et al. (2016), also occur in a similar range of latitudes and altitudes. Beker et al. (2016) reported that H. clavulipes cannot molecularly be separated from H. oreophilum based on five loci. Both are here synonymized with H. palustre. Molecularly, and not only in terms of ITS, H. palustre is very similar to H. fuscatum, H. hygrophilum, and H. nigellum. Since publishing H. fuscatum in 2016, based on just five collections, more intraspecific ITS variation has been observed in H. fuscatum so that the delimitation by ITS data between these two species has become less straight forward.
Geographic distribution
Phenology
  • arrow_drop_downarrow_drop_upAdditional cited collections

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