Hebeloma hiemale Bres.; Fung. trident. 2(11-13): 52, 1892.

Eberhardt, Ursula, Beker, Henry J., Borgen, Torbjorn, Knudsen, Henning, Schuetz, Nicole & Elborne, Steen A., 2021, A survey of Hebeloma (Hymenogastraceae) in Greenland, MycoKeys 79, pp. 17-118 : 17

Hebeloma hiemale Bres.; Fung. trident. 2(11-13): 52, 1892. Fig. 31

Macroscopic description.

Cap 1.1-8.0 cm in diameter, convex to umbonate, often broadly, margin smooth, sometimes involute, undulate in older specimens, tacky when moist, occasionally spotted, rarely hygrophanous, almost uniformly colored or variably bicolored, at center warm buff to grayish buff or dark pinkish buff through honey or ochraceous and dark olive buff to umber and clay pink or orange brown, at margin pale cream to cream to honey or pale pinkish buff, without any traces of veil. Lamellae when young very pale but becoming darker with maturity, emarginate to adnate, maximum depth 3-7 mm, number of lamellae {L} 35-71, droplets usually visible with naked eye or × 10 lens, occasionally absent, white fimbriate edge present. Stem (1.3-)1.5-10.75(-12) × (0.3-)0.35-1.2 {median} × (0.3-)0.55-1.4(-1.5) {base} cm, stem Q 2.5-17.2(-17.4), whitish to pale brownish at middle, hardly discoloring, gradually sordid brown in the mid portion, cylindrical to clavate, rarely subbulbous, usually pruinose particularly at apex, rarely with mycelial chords. Context firm, stem interior stuffed, becoming hollow with age and often with a superior wick, flesh often discoloring from base, particularly after handling. Smell raphanoid, sometimes with hints of cocoa, rarely absent. Taste mild to weakly bitter. Spore color dark olive buff or grayish brown through brownish olive to umber.

Microscopic description.

Spores amygdaloid, limoniform, variably papillate, on ave. 10.0-12.5 × 5.0-7.5 µm, ave. Q = 1.6-2.1, yellow to yellow brown, usually guttulate, weakly, occasionally distinctly ornamented ((O1) O2 (O3)), perispore not or somewhat loosening (P0P1 (P2)), indistinctly to weakly dextrinoid ((D0) D1D2). Basidia 23-39(-43) × 7-10 µm, ave. Q = 3.2-4.3, mostly four-spored, often stipitate. Cheilocystidia usually clavate-lageniform, but occasionally slenderly clavate, clavate-stipitate, cylindrical or ventricose, rarely spathulate-lageniform or tapering, occasionally characteristically with apical or median wall thickening, sometimes geniculate or septate, rarely sinuate or rostrate, on ave. 40-65 × 5.5-9 (apex) × 3.5-5 (middle) × 4.5-7.5 (base) µm, ratios A/M = 1.46-2.48, A/B = 0.97-1.8, B/M = 1.26-1.77. Epicutis an ixocutis, 60-200 µm thick (measured from exsiccata), maximum hyphae width 4-6 µm, sometimes encrusted, trama elements beneath subcutis cylindrical, ellipsoidal, sausage-shaped up to 17 µm wide. Caulocystidia similar to cheilocystidia, up to 75 µm long, often septate, septa sometimes clamped, many cylindrical.

Collections examined.

S-Greenland, Kangilinnguit, 61.21°N, 48.12°W, 20 Aug 2018, H. Knudsen (HK18.269, C-F-111112), 125 m, in tundra. Kangilinnguit, Arsuk Fjord, 61.23°N, 48.07°W, 19 Aug 2018, S.A. Elborne (SAE-2018.225-GR, C-F-112771), 100 m, with Salix glauca and Bistorta vivipara in heathland. Kangilinnguit, at Grønnedal Hut, 61.23°N, 48.08°W, 15 Aug 1985, T. Borgen (TB85.182, C-F-103466), 400 m, with Salix herbacea and Harrimanella hypnoides in snowbed. Nanortalik, 60.08°N, 45.14°W, 12 Aug 1991, T. Borgen (TB91.136, C-F-103465), 50 m, with Salix sp. Nanortalik municipality, Qinngua valley, 60.14°N, 45°W, 9 Aug 1991, T. Borgen (TB91.112, C-F-119782), 230 m, with Salix glauca in scrubland. Narsaq, 60.91°N, 46.05°W, 13 Aug 1993, E. Rald (ER 93.330, C-F-104550), 20 m, with Salix glauca and Betula glandulosa in heathland. Narsaq, 60.91°N, 46.05°W, 3 Aug 1993, E. Rald (ER 93.152, C-F-104318), 20 m. Narsarsuaq, Hospitalsdalen, 61.1715°N, 45.41°W, 10 Aug 1984, T. Læssøe (TL 84.608, C-F-119794), 60 m. Paamiut, 62.01°N, 49.4°W, 8 Aug 1981, T. Borgen (TB81.112, C-F-103552), 10 m, with Salix glauca . Paamiut, 62.01°N, 49.4°W, 27 Aug 1993, T. Borgen (TB93.159, C-F-103496), 20 m, with Salix herbacea . Paamiut, 62.01°N, 49.4°W, 31 Aug 1995, T. Borgen (TB95.114, C-F-103504), 30 m, with Salix glauca , Betula glandulosa in heathland. Paamiut, 62.01°N, 49.4°W, 6 Sep 1990, T. Borgen (TB90.087, C-F-103540), 10 m, with Salix glauca . Paamiut, 62.01°N, 49.4°W, 6 Sep 1990, T. Borgen (TB90.084a, C-F-103549), 10 m, with Salix glauca . Paamiut, 62.01°N, 49.4°W, 8 Sep 1990, T. Borgen (TB90.104a, C-F-103550), 10 m, with Salix glauca and Salix herbacea . Paamiut, 61.99°N, 49.66°W, 22 Aug 2008, T. Borgen (TB08.157, C-F-106752), 15 m, with Salix glauca and Bistorta vivipara . Paamiut, 62.01°N, 49.4°W, 31 Aug 1993, T. Borgen (TB93.183, C-F-103497), 50 m, with Salix glauca in heathland. Paamiut, 62.01°N, 49.4°W, 8 Sep 1993, T. Borgen (TB93.210, C-F-103499), 10 m. Paamiut, 62.01°N, 49.4°W, 1 Sep 1993, T. Borgen (TB93.187, C-F-103498), 10 m, with Salix herbacea . Paamiut, 62.01°N, 49.4°W, 28 Aug 1993, T. Borgen (TB93.155, C-F-103500), 10 m, with Salix herbacea in snowbed. Paamiut, Kangilineq / Kvaneøen, 61.95°N, 49.47°W, 25 Aug 1985, T. Borgen (TB85.250, C-F-119786), 20 m, with Salix arctophila in fenland. Paamiut, Navigation School area, 62.01°N, 49.4°W, 13 Aug 1990, T. Borgen (TB90.032, C-F-103543), 10 m, with Salix herbacea in snowbed. Paamiut, Navigation School area, 62.01°N, 49.4°W, 13 Aug 1990, T. Borgen (TB90.019, C-F-103544), 10 m, with Salix herbacea and Bistorta vivipara . Paamiut, Taartoq/ Mørke Fiord, 61.99°N, 49.66°W, 29 Aug 1998, T. Borgen (TB98.201, C-F-104292), 25 m, with Salix glauca . Qaqortoq, 60.72°N, 46.04°W, 13 Aug 1993, E. Rald (ER 93.302, C-F-104551), 30 m, with Salix glauca . W-Greenland: Kangerlussuaq, 67.01°N, 50.72°W, 11 Aug 1986, T. Borgen (TB86.203, C-F-103568), 50 m. Kangerlussuaq, near a glacier, 67.03°N, 50.64°W, 12 Aug 2000, E. Ohenoja (EO12.8.00.1, OULU F050224), 229 m. Kangerlussuaq, c. 2 km W of the Airport, Mt. Hassel, 67.012°N, 50.856°W, 10 Aug 2000, A-M. Larsen, T. Borgen (TB00.069, C-F-103515), 50 m, with Salix glauca in copse. Kangerluarsunnguaq, Kobbefjord, end of fiord, 64.14°N, 51.35°W, 26 Aug 2018, S.A. Elborne (SAE-2018.357-GR, C-F-112904), 100 m, with Salix glauca in copse. NW below Nasaasaaq, E-valley, E of Sisimiut, 66.93°N, 53.61°W, 18 Aug 2000, E. Horak (ZT8901, ZT8901), 50 m, with Salix glauca . N-Greenland: Daneborg, 0.5 km E of Airstrip, 74.2°N, 20.1°W, 29 Jul 2006, T. Borgen (TB06.033, C-F-119770), 20 m, with Dryas in heathland. Daneborg, slope NW of The Weather Station, 74.2°N, 20.1°W, 31 Jul 2006, T. Borgen (TB06.061, C-F-119769), 20 m, with Dryas sp. in heathland. Zackenberg, 74.5°N, 21°W, 3 Aug 2006, T. Borgen (TB06.081, C-F-119777), 50 m, with Dryas integrifolia and Dryas octopetala in scrubland. Zackenberg, 74.5°N, 21°W, 7 Aug 2006, T. Borgen (TB06.120, C-F-119765), 50 m, with Dryas in heathland. Zackenberg, 74.5°N, 21°W, 2 Aug 2006, T. Borgen (TB06.067, C-F-119743), 50 m, with Dryas sp. and Salix arctica in scrubland. Zackenberg, Aucellabjerg, 74.5°N, 21°W, 27 Jul 1999, T. Borgen (TB99.118, C-F-119756), 150 m, with Dryas sp. and Salix arctica in scrubland. Zackenberg, Aucellabjerg, 74.5°N, 21°W, 20 Aug 2006, T. Borgen (TB06.250, C-F-119764), 400 m, with Dryas sp., Bistorta vivipara and grassland. Zackenberg, Aucellabjerg, 74.5°N, 21°W, 9 Aug 2006, T. Borgen (TB06.128, C-F-119767), 300 m, with Dryas sp. in heathland. Zackenberg, Aucellabjerg, 74.5°N, 21°W, 11 Aug 1999, T. Borgen (TB99.280, C-F-119802), 150 m, tundra. Zackenberg, between West River and Solkæret, 74.5°N, 21°W, 9 Aug 1999, T. Borgen (TB99.258, C-F-119745), 30 m, with Dryas sp. and Salix arctica in scrubland. Zackenberg, just W of Kærelv, 74.5°N, 21°W, 30 Jul 1999, T. Borgen (TB99.160, C-F-119809), 30 m, with Salix arctica and Bistorta vivipara in solifluction lobe in snowbed. Zackenberg, shortly E of Kærelv, 74.5°N, 21°W, 13 Aug 1999, T. Borgen (TB99.304, C-F-119803), 50 m, with Dryas sp. in heathland. Zackenberg, Ulvehøj, 74.5°N, 21°W, 29 Jul 1999, T. Borgen (TB99.146, C-F-104296), 40 m, with Salix arctica in scrubland. E-Greenland: Jameson Land, Nerlerit Inaat/Constable Pynt, delta of Gåseelv valley, 70.76°N, 22.65°W, 8 Aug 2017, T. Borgen (TB17C.078, C-F-106777), 40 m, with Salix glauca in copse. Jameson Land, Nerlerit Inaat/Constable Pynt, Primulaelv, 70.74°N, 22.67°W, 7 Aug 2017, T. Borgen (TB17C.072, C-F-106776), 180 m, with Bistorta vivipara in heathland.

Distribution.

One of the five most commonly recorded Hebeloma species in Greenland, with 11.6% of the records. Common and widespread in Europe; also recorded from other parts of North America (https://mycoportal.org/portal/collections/list.php, accessed 2 Dec 2020). To our knowledge not recorded from alpine sites in Europe ( Beker et al. 2016).

Habitat and ecology.

Forty-four collections, mainly with Salix glauca (15), Dryas (10) and S. herbacea (6). Minor hosts are S. arctophila and S. arctica , and two collections were difficult to separate from the roots of Bistorta vivipara . Found in all types of habitats from scrubland, grassland and heathland to snowbeds. In the Rocky Mountains, the hosts are S. arctica , S. glauca , S. planifolia , S. reticulata , Betula nana and Dryas ( Cripps et al. 2019). For lowland Europe, there is a long list of deciduous trees as probable hosts ( Beker et al. 2016).

Hebeloma sect. Velutipes Vesterh.; Ann. Micol. A. G. M. T. 1: 60, 2004.

Stem velute, usually pruinose at least at apex, bulbous. Cheilocystidia gently clavate towards the apex, occasionally ventricose, spores rather strongly to very strongly dextrinoid.

Figures

Figure 31. Hebeloma hiemale A SAE- 2018.225, photograph S. A. Elborne B distribution of cited collections C spores x 1600 and D cheilocystidia x 1000 of SAE- 2018.225 in Melzer's reagent. Scale bars: 5 µm; microphotographs H. K. J. Beker.