A phylogenetic and morphological study of the genus Dermoloma (Agaricales, Tricholomataceae) in Europe and North America exposes inefficiency of opportunistic species descriptions Author Adamčíková, Katarína 0000-0001-5407-5233 Department of Plant Pathology and Mycology, Institute of Forest Ecology, Slovak Academy of Sciences Zvolen, Akademická 2, 949 01 Nitra, Slovakia Author Kiran, Munazza 0000-0002-3671-9039 Laboratory of Molecular Ecology and Mycology, Institute of Botany, Plant Science and Biodiversity Center, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia & Department of Botany, Division of Science & Technology, University of Education, Lahore, Pakistan Author Caboň, Miroslav 0000-0002-2255-3816 Laboratory of Molecular Ecology and Mycology, Institute of Botany, Plant Science and Biodiversity Center, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Author Matheny, Brandon P. 0000-0003-3857-2189 Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, Tennessee, USA Author Sánchez-García, Marisol 0000-0002-0635-6281 Department of Forest Mycology and Plant Pathology, Swedish University of Agricultural Sciences, Box 7026, 750 07 Uppsala, Sweden Author Arnolds, Eef Department of Forest Ecology, Wageningen University, P. O. Box 9101, 6700 HB Wageningen, Netherlands Author Caboňová, Michaela 0000-0002-2207-6183 Laboratory of Molecular Ecology and Mycology, Institute of Botany, Plant Science and Biodiversity Center, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Author Corriol, Gilles https://orcid.org/0009-0008-7456-9175 Conservatoire botanique national des Pyrénées et de Midi-Pyrénées, Vallon de Salut. B. P. 315, 65203, Bagnères-de-Bigorre Cedex, France Author Dima, Bálint 0000-0003-2099-3903 Department of Plant Anatomy, Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1 / C, H- 1117 Budapest, Hungary Author Friebes, Gernot 0000-0002-0996-3634 Studienzentrum Naturkunde, Universalmuseum Joanneum, Weinzöttlstraße 16, 8045 Graz, Austria Author Griffith, Gareth W. 0000-0001-6914-3745 Department of Life Sciences, Pont Cledwyn, Aberystwyth University, Aberystwyth SY 23 3 DD, Wales, UK Author Grootmyers, Django 0000-0002-1042-4902 Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, Tennessee, USA Author Harries, David 0000-0002-5477-0938 Somerton Cottage, Hundleton, Pembroke, Wales, UK Author Karich, Alexander https://orcid.org/0009-0005-9295-4163 Technische Universität Dresden - Internationales Hochschulinstitut Zittau, Markt 23, 02763 Zittau, Germany Author Mešić, Armin 0000-0002-4479-600X Laboratory for Biological Diversity, Ruđer Bošković Institute, Bijenička cesta 54, HR- 10000 Zagreb, Croatia Author Mihaljevič, Martin 0000-0002-4875-9345 Institute of Geochemistry, Mineralogy and Mineral Resources, Faculty of Science, Charles University, Prague 2, Albertov 6, Czech Republic Author Moreau, Pierre-Arthur 0000-0003-4783-3643 Laboratoire de Génie Civil et géo-Environnement, University of Lille, ULR 4515 – LGCgE, 3 rue du Pr Laguesse, F- 59000 Lille, France Author Pošta, Ana 0000-0003-1715-1609 Laboratory for Biological Diversity, Ruđer Bošković Institute, Bijenička cesta 54, HR- 10000 Zagreb, Croatia Author Shapkin, Vasilii 0000-0002-8394-3009 Laboratory of Molecular Ecology and Mycology, Institute of Botany, Plant Science and Biodiversity Center, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Author Tkalčec, Zdenko 0000-0002-1148-9425 Laboratory for Biological Diversity, Ruđer Bošković Institute, Bijenička cesta 54, HR- 10000 Zagreb, Croatia Author Vizzini, Alfredo 0000-0001-8390-6446 Department of Life Sciences and Systems Biology, University of Torino, Vale P. A. Mattioli 25, 10125 Torino, Italy Author Vondrovicová, Lenka 0000-0003-2791-3160 Institute of Geochemistry, Mineralogy and Mineral Resources, Faculty of Science, Charles University, Prague 2, Albertov 6, Czech Republic Author Adamčík, Slavomir 0000-0003-2156-5767 Laboratory of Molecular Ecology and Mycology, Institute of Botany, Plant Science and Biodiversity Center, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia & Department of Botany, Faculty of Natural Sciences, Comenius University Bratislava, Révová 39, 811 02 Bratislava, Slovakia Author Jančovičová, Soňa 0000-0002-9959-3314 Department of Botany, Faculty of Natural Sciences, Comenius University Bratislava, Révová 39, 811 02 Bratislava, Slovakia text IMA Fungus 2025 2025-07-10 16 e 157337 e 157337 journal article 310963 10.3897/imafungus.16.157337 94e787cd-c3e5-400f-ad11-19727fbfa928 Dermoloma cuneifolium (Fr.) Singer ex Bon , Doc. Mycol. 17 (65): 51. 1986. Figs 20 c – f , 22 Dermoloma emilii-dlouhyi Svrček , Česká Mycol. 20 (3): 147. 1966 . Syn. Neotype . (designated by Arnolds 1992 ): Sweden • Småland , Femsjö , the slope east of Avaberget, amongst mosses , low Hieracium sp. , Trifolium pretense etc., in disused, partly Calluna grown pasture-land , 19 Sep 1948 , S. Lundell and G. Haglund 5101 ( UPS-F-631065 ) . Distinguishing characters. European species; basidiomata medium sized; pilei up to 35 mm in diameter; stipes 2–6 mm wide; lamellae ochraceous-gray to brownish gray; spores inamyloid, 4.7–5.3 × 3.5–4 μm; basidia up to 6.5 μm wide. Pileus (8 –) 12–35 (– 44) mm; convex, soon expanding to plane, often weakly depressed, sometimes indistinctly umbonate, sometimes lobate; margin usually not striate, indistinctly translucently striate when wet; surface near margin smooth, near center radially rugulose and veined, sometimes pitted or rough, not hygrophanous; color when young dark brown (6 F 2, 6 F 3, 6 F 4, 7 F 3, 7 F 4), when mature near margin usually grayish brown (5 D 3, 6 D 3,6E 3,7E 3), often brownish gray (5 D 2, 6 C 2), brown (5 E 4,6E 4,7E 4) to dark brown (6 F 3, 6 F 4, 7 F 3), rarely grayish ochraceous (5 B 3), brownish ochraceous (5 C 3) or dark blond (5 D 4), towards margin often gradually passing to narrow paler brownish gray (6 C 2), ochraceous-gray (5 B 2) to almost white zone, near center usually dark brown (6 F 2, 6 F 3, 6 F 4, 7 F 3, 7 F 4, 7 F 5) to black, sometimes brown (5 E 4,6E 4,7E 4), rarely grayish brown (6 E 3). Stipe (17 –) 21–50 (– 55) × (1 –) 2–6 (– 8) mm; cylindrical or fusiform, usually narrowed towards the base, flexuous; surface near lamellae pruinose or granulose, towards the base granulose or finely fibrillose, finely longitudinally striate; color near lamellae white, pale gray (B 1), ochraceous-gray (5 B 2), brownish gray (5 C 2, 6 C 2), near the base usually brownish gray (5 C 2, 5 D 2, 6 C 2, 6 D 2) or grayish brown (5 D 3,5E 3, 6 D 3), sometimes ochraceous-gray (5 B 2, 6 B 2), rarely grayish ochraceous (5 B 3), brownish ochraceous (5 C 3) or brown (5 E 4). Lamellae L = (19 –) 23–38 (– 42), l = 1–7; 4–9 mm wide; adnate-emarginate, sometimes decurrent with tooth or adnexed; color usually ochraceous-gray (5 B 2) or brownish gray (5 C 2, 6 C 2), often almost white, rarely grayish brown (6 D 3); edges entire, rarely serrulate. Context fragile; odor farinaceous. Dermoloma cuneifolium ( UPS-F-631065 , neotype), microscopic elements. a Spores; b caulocystidia; c basidia and basidioles; d marginal cells; e pileipellis elements near the pileus margin; f pileipellis elements near the pileus center. Scale bar: 5 µm for spores and 10 µm for other elements. Spores (4.2 –) 4.7–5 – 5.3 (– 6.1) × (3.2 –) 3.5–3.8 – 4 (– 4.4) μm; broadly ellipsoid to ellipsoid, Q = (1.14 –) 1.24–1.32 – 1.40 (– 1.53); walls inamyloid, sometimes thick-walled and dextrinoid; hilar appendage 0.5–1 μm long. Basidia (20 –) 23.5–25.9 – 28.5 (– 32) × (5 –) 5.5–6 – 6.5 (– 7) μm; clavate; with 4 sterigmata. Basidioles first cylindrical, then clavate, ca. 3–6.5 μm wide. Marginal cells (10 –) 14–19.1 – 24 (– 28) × (2 –) 3.5–4.7 – 6 (– 8) μm; cylindrical or clavate. Pileipellis 55–65 μm deep; suprapellis of mainly one, occasionally two layers of inflated cells, gradually passing to 18–25 μm deep subpellis of densely packed, irregularly oriented, puzzled, 3–8 μm wide hyphae, not sharply delimited from horizontally oriented hyphae in trama; hyphal terminations with brownish parietal pigments, thin-walled or occasionally thickened up to 1 μm and with brown incrusted pigments especially near septa of terminal cells. Terminal cells near pileus margin (11 –) 28–37.4 – 46.5 (– 68) × (6 –) 13.5–17.9 – 22 (– 33) μm; usually sphaeropedunculate, sometimes obpyriform or clavate; subterminal cells usually narrower and implemented in subpellis, often branched, sometimes ventricose and short, rarely with lateral swellings. Terminal cells near pileus center (13 –) 27–37 – 47.5 (– 59) × (5.5 –) 11–16.4 – 21.5 (– 32) μm; usually obpyriform, clavate or sphaeropedunculate, rarely ellipsoid or clavate-lageniform, sometimes with one or two central constrictions; subterminal cells similar to cells near margin. Caulocystidia (10 –) 25–37.6 – 50.5 (– 81) × (3 –) 4–6.6 – 9 (– 12) μm; clavate or cylindrical, usually not or only slightly flexuous, often clustered in small ascending fascicules, sometimes individual and repent; occasionally with slightly thickened walls up to 0.5 μm, often with crystalline or granulose yellow incrustations. Clamp connections present. Distribution and ecology. Probably the most common Dermoloma species, widely distributed in Europe; in semi-natural grasslands, dunes, forests, cemeteries and parks, often collected in habitats with low conservation value as the single Dermoloma species. Additional material studied. Austria • Burgenland , Oberwart , Rechnitz , Galgenberg , elev. 344 m , coord. 47°17'52"N , 16°25'09"E , semi-dry grassland, soil among mosses and grass , 17 Nov 2019 , G. Friebes GF 20190133 ( SAV F-23422 ) ; • Steiermark , Südoststeiermark , Straden , Stradner Kogel , Berghölzer , elev. 411 m , coord. 46°49'53"N , 15°55'02"E , soil among mosses and grass , 6 Oct 2019 , G. Friebes GF 20190080 ( SAV F-23426 ) . Croatia • Gorski kotar area , Crni Lug – Velika Voda , coord. 45°24'36"N , 14°42'15"E , mowed grassland with short grass and mosses , 4 Oct 1998 , A. Mešić ( CNF 5/204 ) ; • 2.2 km SE / S-SE of Brlog Ozaljski village , near Ozalj , coord. 45°36'33"N , 15°25'11"E , mowed grassland with short grass and mosses, Calluna vulgaris , Juniperus communis , Pteridium aquilinum , 16 Oct 2010 , Z. Tkalčec and I. Tkalčec ( CNF 1/6057 ) . Czech Republic • Brdské hřebeny Mts. , Vižina , mossy forest meadow , 28 Sep 1965 , E. Dlouhý ( PRM 610931 , holotype of D. emilii-dlouhyi ) . France • Aveyron , Le Fel , Le Mas , coord. 44°39'27"N , 02°30'55"E , old grassland , 15 Nov 2016 , C. Hannoire CH 16111509 ( BBF , as D. fuscobrunneum ) ; • Corse du Sud , Bastelica , pont de Zipitoli , coord. 41°57'38"N , 09°00'08"E , Quercus ilex thicket on mineral-rich clay soil , 19 Nov 2014 , P. - A. Moreau PAM 14111900 ( LIP , as D. cf. atrocinereum ) ; • Hautes-Pyrénées , Payolle , coord. 42°55'57"N , 00°16'03"E , acidophilic grazed grassland ( Nardion ) , 15 Oct 2016 , G. Corriol GC 16101502 ( BBF , as D. fuscobrunneum ) ; • Hautes-Pyrénées , Tramezaïgues , Fontaine des Usclats , coord. 42°47'32"N , 00°17'26"E , grazed grassland , 24 Sep 2018 , C. Hannoire CH 18092415 ( BBF , as D. fuscobrunneum ) ; • Pas-de-Calais , Merlimont , Réserve biologique , coord. 50°26'52"N , 01°35'10"E , fixed dune, open grassland on sand , 10 Nov 2014 , P. - A. Moreau PAM 14111009 ( LIP , as D. cf. pseudocuneifolium ) . Germany • Gottesacker , Herrnhut , coord. 51°01'11"N , 14°44'53"E , ancient graveyard , 25 Aug 2016 , A. Karich and R. Ullrich IHI- 16 Der 01 ( GLM-F 137754 ) ; • Niedersachsen , Meppen , Mepper Kuhweide , poor grassland on dry calcareous sand , 5 Nov 2015 , E. Arnolds Arnolds 15-95 ( L , as D. phaeopodium ) ; • Schleswig-Holstein , Winderatter See , 11 Nov 2015 , T. Böhning AG 110 ( ELTE ) . Italy • Marche , Cantiano (PU) , Bosco di Tecchie , 17 Oct 2012 , G. Consiglio , M. Maletti and L. Polidori GC 12181 ( AMB 15102 ) ; • Sardinia , Tempio Pausania , Monti Avoni , 28 Dec 2008 , M. Contu 28. XII. 2008 ( AQUI , as D. fuscobrunneum ) ; • Savona , Sassello , Periaschi , 2 Nov 2013 , F. Bocianolo G 3180 ( GDOR ) . Romania • Cindrel Mts. , pasture, 0.5 km SW of Poiana Sibiului , elev. 940–950 m , 45°47'52"N , 23°42'38"E , terrestrial , 26 Sep 2015 , S. Jančovičová ( SAV F-4702 ) ; • Gilau Mts. , pasture 750 m E of Someşu Rece , elev. 500–530 m , coord. 46°43'21"N , 23°22'20"E , terrestrial , 4 Oct 2014 , S. Adamčík ( SAV F-4265 ) . Slovakia • Považský Inovec Mts. , 2 km SE of Kálnica , coord. 48°45'18"N , 17°56'55"E , elev. 300 m , terrestrial in semi-natural grassland , 24 Oct 2014 , S. Jančovičová ( SLO 674 ) ; • ibid., 24 Oct 2014 , S. Jančovičová ( SLO 675 ) ; • ibid., 24 Oct 2014 , S. Jančovičová ( SLO 676 ) ; • ibid., 24 Oct 2014 , S. Jančovičová ( SLO 677 ) ; • ibid., 24 Oct 2014 , S. Jančovičová ( SLO 678 ) ; • ibid., 24 Oct 2014 , S. Jančovičová ( SLO 679 ) . Sweden • Småland , Femsjö par. , “ Haggårds tomter ”, NW of Skattegårten , 6 Sep 1948 , S. Lundell and G. Haglund ( UPS-F-631060 ) ; • Småland , Femsjö par. , Källebo , mossy grassland , 4 Sep 1948 , S. Lundell and G. Haglund ( UPS-F-631061 ) ; • Småland , Femsjö par. , the slope east of Arvaberget towards Artvamaden , pasture, among Hieracium pilosella , 20 Sep 1948 , S. Lundell and G. Haglund ( UPS-F-631063 ) ; • Småland , Femsjö par. , Femsjö village , Rysslandsåkern , mossy grassland , 13 Sep 1948 , S. Lundell and G. Haglund ( UPS-F-631064 ) . United Kingdom • England , Shropshire , Shrewsbury cemetery , coord. 52°41'46"N , 02°45'32"E , terrestrial in lawn , 21 Oct 2014 , S. Adamčík ( SAV F-4361 ) ; • Wales , Dugoed , 2 Nov 2004 , G. W. Griffith SH 804522 ( ABS ) ; • Wales , Mynachdy , Brignant grassland , terrestrial , 26 Oct 2016 , G. W. Griffith ( SAV F-20076 ) ; • Wales , Montgomery , Newtown cemetery , coord. 52°31'06"N , 03°18'06"E , terrestrial in lawn , 24 Oct 2014 , P. David ( SAV F-4393 ) ; • ibid., 24 Oct 2014 , D. Harries ( SAV F-4394 ) ; • ibid., 24 Oct 2014 , D. Harries ( SAV F-4395 ) ; • Wales , Powys , Gregynog grounds , coord. 52°34'04"N , 03°21'06"E , terrestrial in lawn , 24 Oct 2014 , D. Harries ( SAV F-4396 ) ; • ibid., 24 Oct 2014 , D. Harries ( SAV F-4397 ) ; • Wales , Powys , Hay Common , coord. 52°02'34"N , 03°06'19"E , terrestrial in semi-natural grassland , 28 Oct 2014 , S. Adamčík ( SAV F-4435 ) ; • ibid., 28 Oct 2014 , D. Harries ( SAV F-4436 ) ; • ibid., 28 Oct 2014 , D. Harries ( SAV F-4437 ) ; • ibid., 28 Oct 2014 , D. Harries ( SAV F-4438 ) ; • Wales , Pembrokeshire , Somerton farm , coord. 51°39'50"N , 04°59'29"E , terrestrial in pasture , 25 Oct 2014 , S. Adamčík ( SAV F-4407 ) ; • ibid., 25 Oct 2014 , D. Harries ( SAV F-4408 ) ; • Wales , Pembrokeshire , Tufton cemetery , coord. 51°55'02"N , 04°51'07"E , terrestrial in lawn , 27 Oct 2014 , S. Adamčík ( SAV F-4423 ) ; • ibid., 27 Oct 2014 , leg. S. Adamčík ( SAV F-4425 ) . Notes. Dermoloma cuneifolium has inamyloid spores as a member of D. subgenus Dermoloma , section Dermoloma . The section forms a crown clade in the Dermoloma phylogeny with short branch lengths at species rank clades indicating short genetic distances (Fig. 2 ). Morphologically, D. cuneifolium exhibits intermediate characters within this group of very similar species. It has intermediate basidiomata sizes, spores which are relatively small but similar like in several other related species. It is similar, especially to D. simile , D. huartii and D. intermedium . Here, and in similar cases, we strongly suggest to use the key for identification, in combination with the heatmap diagram (Fig. 4 ), chromatogram (Fig. 6 ), plot diagrams (Figs 5 – 7 ) and other supplementary files (Suppl. materials 6 , 7 , 9 ) for the best score when combining multiple morphological characters. We believe that the probability of correct identification of this species may be high, but for precise identification we recommend to use sequences of the ITS region. The sister species to D. cuneifolium is the North American D. fumosidiscum (Fig. 2 ), which is distinctly smaller and has strongly rugulose pileus center. This species was included in the phylogenetic study by Sánchez-García et al. (2021) and the name was assigned by the position of the neotype sequence. It is the type species of Dermoloma .