Hebeloma cistophilum (Photo: H. J. Beker)

Taxonomy

Types: ALGERIA: Boufarik (approx. 36.54°N, 2.87°E, alt. approx. 100 m a.s.l.) under Cistus sp., 1 Dec. 1912, Duvernoy, det: R. Maire (1113) (Lectotype. held at herbarium MPU, HJB1000102). Lectotype designated by Grilli, Maire J.-C., Moreau P.-A. Robich G. (Eds.), Compléments a la Flore des Chamignons Superieurs de Maroc de G. Malençon et R. Bertault. Confédération Européenne de Mycologie Méditerranéenne, Nice: 299-318. : (2009) page 300.

• arrow_drop_downarrow_drop_upType notes
  The lectotype designated by Grilli supersedes the neotype designated by M. Heykoop & F. Esteve-Raventós, Mycotaxon 61: 212 (1997)

Homotypic synonyms:

• Hebelomatis cistophilum (Maire) Locq., Flore Mycologique Vol III - Text. Cortinariales A: 129 (1979) ["1977"]

• arrow_drop_downarrow_drop_upEtymology
  From the plant Cistus and philus– loving (Greek), to emphasise the relationship between this taxon and Cistaceae, with which it is always in ectomycorrhizal association.

• arrow_drop_downarrow_drop_upOriginal diagnosis
  Carpophora gregaria, saepe circinantia, haud hygrophana; sapor mitis l. vix amarescens, odor rapae; caro ex albido fulvescens demum in imo stipite fulva l. subfuscescens; spore in cumulo sordide fusco-ferrugineae (K: 129-130-133). Stipes subaequalis, 2–4 cm x 3–5mm., fibroso-carnosus, e farcto cavus, cum pileo confluens, siccus, sub cortina floccoso-fibrillosus, supra cortinam fibrilloso-striatus, apex pruinosus, albidus, demum fulvescens et base interdum fulvo-fuscescens. Cortina albida, stipitem diu cingulans, mox e sporis sordide fusco-ferruginea. Velum universale in margine pilei haud persistens. Pileus 1–3 cm diam. e convexo l campanulato-convexo applanatus, plus minusve umbonatus, tenuis, carnosus, ochraceo-fulva (M: 142), in disco castanea (K: 114), margine incurvo, laevi, concolore, primum cortina fimbriato, dein glabro. Lamellae confertae, tenues, latae, ventricosae, utrinque attenuatae, cum hymenophoro confluentes, plus minusve adnatae, emarginatae, ex albido fulvae l fulvo-umbrinae, acie floccose albida, haud intervenatae. Lamellulae plus minusve rotundatae, interdum subemarginatae. Lamellarum acies pilis cylindraceis flexuosis heteromorpha, mediostratus regularis; subhymenium tenue ramosum; cystidia nulla; basidia cylindaceo clavata, 4-spora, 32–38 x 8 μm; sporae amygdaliformes, sub lente mellae, 10–11.5 x 6–7 μm, episporio undique aequali subtillime verrucose (verrucis in lacto-phenolis object. immers. ope tantum conspicuis. G -, NH3 -. Hab. in cistetis, rarius in quercetis Mauretaniae, autumn et hieme copioso.

• arrow_drop_downarrow_drop_upEnglish translation
  Basidiomes gregarious and often in rings, not hygrophanous; mild taste later slightly bitter; smell raphanoid, context white then turning brown, in base of stipe becoming reddish brown; spores in mass sordid brown-rusty brown. Stipe subequal, 2–4 cm × 3–5 mm, fibrous-fleshy, solid then hollow, confluent with pileus, dry, under cortina floccose-fibrillose, above cortina fibrillose-striate, apex pruinose, white, then turning yellow brown and base sometimes reddish brown. Cortina white, on stipe in persistent girdles, later turning brown by ripe spores. Velum universale on margin of pileus not persistent. Pileus 1–3 cm diam., convex then campanulate-convex applanate, more or less umbonate, thin-fleshed, very fragile, cuticule more or less peeling, viscid, glabrous. Lamellae crowded, thin, broad, ventricose, attenuate at both sides, confluent with hymenophore, more or less adnate, emarginate, white then reddish brown to umber-brown, with white floccose edge, not interveined. Lamellae more or less rounded, sometimes submarginate. Lamella edge heteromorphous with cylindrical, flexuous hairs; cystidia absent; basidia cylindrical, clavate, four-spored, 32–38 × 8 μm; spores amygdaloid, under lens honey-coloured, 10–11.5 × 6–7 μm, with regular, subtly verrucose episporium (warts in lacto-phenol under oil immersion equally conspicuous). Guaiac and ammonia negative. In Cistus vegetations, rarely in Quercus forest in Mauretania, abundant in autumn and winter.

Description

Description of Hebeloma cistophilum based on 94 collections

• arrow_drop_downarrow_drop_upMacroscopic description
  Pileus: (10) 19–40 (47) mm diameter; shape often convex, occasionally strongly umbonate, umbonate or broadly umbonate, rarely papillate; characters occasionally remains of universal veil, rarely hygrophanous or pruinose; margin characters often smooth, occasionally involute, rarely crenulate, eroded or wavy; viscosity tacky when moist; colour variation often two color, occasionally unicolour; colour at centre occasionally dark brick or yellowish brown, rarely umber, clay-red, ochraceous, orange-brown, cinnamon, sepia, greyish brown or dark fawn.

  Lamellae: attachment usually emarginate, rarely adnate or decurrent tooth; maximum depth 2–5 mm; number of complete lamellae 37–63; presence of tears absent; white fimbriate edge often weak, occasionally present or absent.

  Cortina presence: yes.

  Stipe: (18) 22–55 (70) x 3–9 (12) {median} x 3–8 (12) {basal} mm; stipe Q 2.0–15.0; base shape usually cylindrical, occasionally clavate, rarely tapering or sand bulb; floccosity often fibrillose, occasionally pruinose at apex or floccose at apex, rarely floccose or none; rooting no; thick rhizoids at base absent;

  Context: Texture firm; stipe interior usually hollow, occasionally stuffed, rarely superior wick; stipe flesh discolouring variable occasionally weak; slenderness measure 1.9–21.6; smell often raphanoid, rarely weakly raphanoid, odourless, cocoa, earthy or strongly raphanoid; taste often bitter, occasionally weakly bitter, rarely mild, raphanoid or weakly raphanoid where recorded.

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

  Exsiccata characters: rarely fragile, pale, pileus blackening, rich brown color or stipe blackening.

• arrow_drop_downarrow_drop_upMicroscopic description
  Spores: shape amygdaloid, usually ellipsoid, often ovoid, rarely cylindrical; colour in microscope often yellow, occasionally brown pale or grey yellow, rarely yellow pale; guttules usually yes, rarely no or weak. papilla usually no, rarely weak; Spore Code: O1 O2; P0; D0 D1.

  Basidia: 26–38 (40) x 6–10 μm; ave. Q 3.6–4.9; spore arrangement Not recorded;

  Cheilocystidia: main shape ventricose, usually lageniform; special features observed often septa, rarely geniculate, apical thickening, clamped septa, many collapsed in exsiccata, median thickening or slender; cheilocystidia ratios: A/M = 0.91–1.21; A/B = 0.37–0.74; B/M = 1.60–2.67.

  Pleurocystidia: variable rarely only close to lamella edge.

  Ixocutis: epicutis thickness (measured from exsiccata) up to 175 μm; ixocutis hyphae width up to 7 μm; ixocutis hyphae encrustation yes; shape of trama elements beneath subcutis often thickly sausage-shaped, occasionally ellipsoid, isodiametric, cylindrical, oblong or pyriform up to 12 μm wide.

  Caulocystidia: Similar to cheilocystidia but larger, up to 100 μm.

• arrow_drop_downarrow_drop_upSpore measurements

• arrow_drop_downarrow_drop_upCheilocystidia measurements

• arrow_drop_downarrow_drop_upHabitat and distribution
  Hebeloma cistophilum's preferred habitat appears to be maquis or mixed woodland with decomposed litter. Where only one possible associate was recorded, the most commonly recorded associate was Cistus (81.5%) but Pinus (11.1%), Helianthemum (1.9%), Quercus (1.9%), Salix (1.9%) and Abies (1.9%) were also recorded. In these cases the most commonly recorded families were Cistaceae (83.9%) and Pinaceae (12.5%). We have additional records where Cedrus was recorded as a possible associate, but for these collections a number of possible associates were mentioned. Overall the most commonly recorded families are Cistaceae (89.7%), Pinaceae (33.3%) and Fagaceae (14.9%) The growth habit of our collections was often scattered, occasionally gregarious or caespitose and rarely solitary.

  According to our current data, the species is found on multiple continents with collections found in Europe (66.3%), Temperate Asia (30.4%) and Africa (3.3%). On these continents, collections have been found only in the mediterranean forests, woodlands & scrub WWF biome 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. (Cyprus Mediterranean forests (28.7%), Tyrrhenian-Adriatic sclerophyllous and mixed forests (16.0%) and Iberian sclerophyllous and semi-deciduous forests (10.6%) ecoregions). From collector information, it appears collections have been found in the 1.4 Forest – Temperate (42.9%), 3.8 Shrubland – Mediterranean-type shrubby vegetation (41.3%) and 13.3 Coastal Sand Dunes (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 Southwest (Spain, France, Portugal and Italy) and the Southeast (Italy). Specimens have been collected from 37.0°N to 49.5°N.

  Within Temperate Asia all our records are from Western Asia (Cyprus and Turkey).

  Within Africa we have records from Northern Africa (Algeria) and Macaronesia (Spain).

• arrow_drop_downarrow_drop_upMolecular results
  Hebeloma cistophilum is very well supported by bootstrap in all single locus results, apart from those of mitochondrial loci, as well as in the five locus result. Intraspecifically sequences of H. cistophilum are quite variable but there are no consistent patterns between the results of different loci. Thus there is no suggestion of the presence of hidden species. Unfortunately, but predictably, we have not been able to amplify the ITS or any other locus from the lectotype, collected in 1912, of this species. We are not aware of any ITS sequences from outside Europe that are likely to belong to H. cistophilum. However, there is a sequence from Spain (JQ975961) that was allegedly amplified from a Pinus pinaster root tip. This does suggest that mycorrhizal association with hosts other than Cistaceae is possible, at least if Cistus spp. are present as was reported in this case Rincón et al. 2014).

• arrow_drop_downarrow_drop_upCommentary
  With the persistent presence of a cortina and the lageniform or ventricose cheilocystidia, this taxon clearly belongs in H. sect. Hebeloma. Within this section, it is unusual in that its spore shape is a mixture of ovoid, amygdaloid and ellipsoid. It is in mycorrhizal association with Cistaceae and appears to be the only Hebeloma sp. from this section which has been collected in association with Cistaceae. (We have one unconfirmed record of H. mesophaeum with Helianthemum.) Hebeloma cistophilum frequently has pleurocystidia, which is unusual within the European taxa of this genus. However, we have not always been able to find pleurocystidia and we suspect that they are not always present. When we have found them they are usually sparse. Hebeloma cistophilum can vary a great deal macroscopically. Indeed we have had collections that we believed, in the field, were a different taxon. However, despite their very different macroscopic appearance, we found no consistent microscopic or molecular difference that could indicate further splits. We have therefore continued to treat this as a single taxon, although future researchers may well find consistent differences and grounds on which to split this taxon.

Geographic distribution

Phenology

• arrow_drop_downarrow_drop_upAdditional cited collections

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