Abstract
The ectomycorrhiza forming basidiomycete Hygrophorus penarius sensu lato was recognized as two species, H. penarioides and H. penarius in 2007 based on molecular and morphological evidence. However, this was based on few collections of restricted geographical origin and no studies have performed molecular analysis on additional specimens to date.
To this end we sequenced the nrDNA ITS regions of 29 H. penarius s. l. collections from Hungary and Slovakia and analysed with collections from previous studies. Bayesian and Maximum Likelihood phylogenetic analyses confirmed the separation of H. penarioides and H. penarius in two clades with the majority of the specimens belonging to H. penarioides and only three to H. penarius. Despite the strong support of the lineages, there was no coherent difference in basidiome and spore traits. We also provide the first evidence of an intraspecific nrDNA sequence variation in the ITS2 region of H. penarioides together with morphological variability that has not been reported to date.
On the basis of our result, we conclude that the Quercus associated H. penarioides is far more common in Hungary than the Fagus associated H. penarius and we suggest that identification of the two species in the absence of DNA sequence analysis should be handled with caution, especially when herbarium specimens are examined.
References
Arnolds, E. (1986) Notes on Hygrophoraceae–VII. On the taxonomy and nomenclature of some species of Hygrophorus. Persoonia 13: 69–76.
Arnolds, E. (1990) Tribus Hygrophoreae. In: Bas, C., Kuyper, Th.W., Noordeloos, M.E. & Vellinga, E.C. (Eds.) Flora Agaricina Neerlandica 2. A. A. Balkema, Rotterdam, pp. 115–133.
Babos, M. (1989) The Agaricales s. l. taxa of Hungary. Mikológiai Közlemények, Clusiana 28: 3–234.
Bataille, F. (1910) Flore monographique des hygrophores. Mémoires de la Société d’émulation du Doubs 8 (4): 132–189.
Becker, G. (1954) Un nouvel Hygrophore: Hygrophorus (Limacium) barbatulus. Schweizerische Zeitschrift für Pilzkunde 32: 91–92.
Bohus, G. (1963) New suggestions for preparing fleshy fungi for the herbarium. Mycologia 55: 128–130.
https://doi.org/10.2307/3756389
Bon, M. (1974) Hygrophores du Centre-Est de la France étudiés du salon du Museum 1971. Bulletin mensuel de la Société linnéenne de Lyon 43: 333–343.
Bon, M. (1977) Clé analytique des Hygrophoraceae (suite). Le genre Hygrophorus Fr. Documents Mycologique 7: 25–53.
Bon, M. (1989) Nouveaux taxons (Hygrophoraceae). Documents Mycologique 75: 55–56.
Bon, M. (1992) Die Großpilzflora von Europa. 1. Hygrophoraceae. Übersetzt und bearbeitet von A. Einhellinger, IHW-Verlag, Eching.
Borchsenius, F. (2009) FastGap 1.2. Department of Bio-sciences, Aarhus University, Denmark. Available from: <a href="http://www.aubot.dk/FastGap_home.htm"><span style="text-decoration: underline;">http://www.aubot.dk/FastGap_home.htm</span></a> (accessed 14 February 2019)
Brock, P.M., Doring, H. & Bidartondo, M.I. (2009) How to know unknown fungi: the role of a herbarium. New Phytologist 181: 719–724.
https://doi.org/10.1111/j.1469-8137.2008.02703.x
Bulliard, J.B.F. (1780) Herbier de la France. 2. Chez l’auteur, Didot, Debure, Paris, 48 pp.
Candusso, M. (1997) Hygrophorus s. l. In: Fungi Europaei 6. Libreria Basso, Alassio.
di Marino, E., Montecchio, L. & Agerer, R. (2008) Hygrophorus penarius L. + Fagus sylvatica L. Descriptions of Ectomycorrhizae 11–12: 77–82.
Fernández-Vicente, J., Hidalgo, F. & Oyarzabal, M. (2015) Adiciones al Parque Natural del Gorbea y aledaños, Araba-Bizkaia, País Vasco. Agaricales, Hygrophoraceae (Basidiomycota). Yesca 27: 31–52.
Fries, E.M. (1836a) Corpus Florarum provincialium suecicae 1. Floram Scanicam, Uppsala.
Fries, E.M. (1836b) Anteckningar öfver de i Sverige växande ätliga Svampar. Palmblad, Sebell, Uppsala.
Fries, E.M. (1838) Epicrisis systematis mycologici seu synopsis Hymenomycetum. Uppsala.
Gardes, M. & Bruns, T.D. (1993) ITS primers with enhanced specificity for basidiomycetes–application to the identification of mycorrhizae and rusts. Molecular Ecology 2 (2):113–118.
https://doi.org/10.1111/j.1365-294X.1993.tb00005.x
Gastwirth, L.J., Gel, R.Y., Hui, W.L.W., Lyubchich, V., Miao, W. & Noguchi, K. (2017) lawstat: Tools for Biostatistics, Public Policy, and Law. R package version 3.1. Available from: https://CRAN.R-project.org/package=lawstat (accessed 14 February 2019)
Gouy, M., Guindon, S. & Gascuel, O. (2010) SeaView version 4: a multiplatform graphical user interface for sequence alignment and phylogenetic tree building. Molecular Biology and Evolution 27: 221–224.
https://doi.org/10.1093/molbev/msp259
Hawksworth, D.L. (2013) The oldest sequenced fungal specimen. Lichenologist 45: 131–132.
https://doi.org/10.1017/S0024282912000710
Heim, R. (1948) Deux Hygrophores nouveaux du Jura. Bulletin de la Société Mycologique de France 63: 127–132.
Hesler, L.R. & Smith, A.H. (1963) North American species of Hygrophorus. University of Tennessee Press, Knoxville.
https://doi.org/10.5962/bhl.title.61976
Holec, J. & Wild, J. (2011) Fungal diversity in sandstone gorges of the Bohemian Switzerland National Park (Czech Republic). Czech Mycology 63 (2): 243–263.
Jacobsson, S. & Larsson, E. (2007) Hygrophorus penarioides, a new species identified using morphology and ITS sequence data. Mycotaxon 99: 337–343.
Kalmár, Z. (1966) The frequency of the higher mushroom species in Hungary. Mikológiai Közlemények, Clusiana 4: 17–26.
Kirk, P.M., Cannon, P.F., Minter, D.W. & Stalpers, J.A. (2008) Ainsworth & Bisby’s dictionary of the fungi. 10th edition. CABI International, Wallingford.
https://doi.org/10.1079/9780851998268.0000
Konrad, P. & Maublanc, A. (1937) Icones selectae fungorum, vol 6., Paul Lechevalier, Paris.
Kovalenko, A. (1999) The arctic-subarctic and alpine-subalpine component of the Hygrophoraceae in Russia. Kew Bulletin 54: 695–704.
https://doi.org/10.2307/4110865
Kovalenko, A. (2012) Hygrophorus Fr. In: Knudsen, H. & Vesterholt, J. (Eds.) Funga Nordica. Agaricoid, boletoid, clavarioid, cyphelloid and gastroid genera. Nordsvamp, Copenhagen, pp. 282–293.
Krieglsteiner, G.J. (Ed.) (2000) Die Großpilze Baden-Württembergs. Band 1. Ulmer Verlag, Stuttgart.
Krieglsteiner, G.J. (Ed.) (2001) Die Großpilze Baden-Württembergs. Band 3. Ulmer Verlag, Pliezhausen.
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33 (7): 1870–1874.
https://doi.org/10.1093/molbev/msw054
Larsson, E. & Jacobsson, S. (2004) Controversy over Hygrophorus cossus settled using ITS sequence data from 200 year-old type material. Mycological Research 108: 781–786.
https://doi.org/10.1017/S0953756204000310
Lodge, D.L., Padamsee, M., Matheny, P.B., Aime, M.C., Cantrell, S.A., Boertmann, D., Kovalenko, A., Vizzini, A., Dentinger, B., Kirk, P., Ainsworth, A.M., Moncalvo, J.-M., Vilgalys, R., Larsson, E., Lücking, R., Griffith, G.W., Smith, M.E., Norvell, L.L., Desjardin, D.E., Redhead, S.A., Ovrebo, C.L., Lickey, E.B., Ercole, E., Hughes, K.W., Courtecuisse, R., Young, A., Binder, M., Minnis, A.M., Lindner, D.L., Ortiz-Santana, B., Haight, J., Læssøe, T., Baroni, T.J., Geml, J. & Hattori, T. (2014) Molecular phylogeny, morphology, pigment chemistry and ecology in Hygrophoraceae (Agaricales). Fungal Diversity 64: 1–99.
https://doi.org/10.1007/s13225-013-0259-0
Lowy, B. (1958) On preparing fleshy fungi for the herbarium. Mycologia 50: 442–444.
https://doi.org/10.2307/3756081
Matheny, P.B., Curtis, J.M., Hofstetter, V., Aime, M.C., Moncalvo, J.M., Ge, Z.W., Slot, J.C., Ammirati, J.F., Baroni, T.J., Bougher, N.L., Hughes, K.W., Lodge, D.J., Kerrigan, R.W., Seidl, M.T., Aanen, D.K., DeNitis, M., Daniele, G.M., Desjardin, D.E., Kropp, B.R., Norvell, L.L., Parker, A., Vellinga, E.C., Vilgalys, R. & Hibbett, D.S. (2006) Major clades of Agaricales: a multilocus phylogenetic overview. Mycologia 98: 982–995.
https://doi.org/10.1080/15572536.2006.11832627
Melis, J.L.L., Mir, G. & Prats, M.C. (2017) Aportación Al Catálogo Micológico De Las Illes Balears. Menorca, Ii. Micobotánica-Jaén Año XII Nº1 Enero-Marzo 2017: 66–99.
Nagy, L.G., Kocsubé, S., Csanádi, Z., Kovács, G.M., Petkovits, T., Vágvölgyi, Cs. & Papp, T. (2012) Re-mind the gap! Insertion-deletion data reveal neglected phylogenetic potential of the nuclear ribosomal internal transcribed spacer (ITS) of fungi. PLoS One 7:e49794.
https://doi.org/10.1371/journal.pone.0049794
Osmundson, T.W., Robert, V.A., Schoch, C.L., Baker, L.J., Smith, A., Robich, G., Mizzan, L. & Garbelotto, M.M. (2013) Filling gaps in biodiversity knowledge for macrofungi: contributions and assessment of an herbarium collection DNA barcode sequencing project. PLoS One 8: e62419.
https://doi.org/10.1371/journal.pone.0062419
Peck, C.H. (1898) New species of fungi. Bulletin of the Torrey Botanical Club 25 (6): 321–328.
https://doi.org/10.2307/2478129
Pecoraro, L., Perini, C., Salerni, E. & Dominicis, V.D. (2007) Contribution to the knowledge of the mycological flora of the Pigelleto Nature Reserve, Mt. Amiata (Italy). Flora Mediterranea 17: 143–163.
R Core Team (2015) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.Available from: https://www.R-project.org/ (accessed 14 February 2019)
Rimóczi, I., Siller, I., Vasas, G., Albert, L., Vetter, J. & Bratek, Z. (1999) Magyarország nagygombáinak javasolt vörös listája. Mikológiai Közlemények, Clusiana 38 (1–3): 107–132.
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574.
https://doi.org/10.1093/bioinformatics/btg180
R Studio Team (2015) RStudio: Integrated Development for R. RStudio Inc., Boston, MA. Available from: http://www.rstudio.com/ (accessed 14 February 2019)
Rücker, T. (1990) Die Pilzflora der Gaisbergwälder bei Salzburg, Österreich. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Österreich 127: 165–183.
Seitzman, B.H., Ouimette, A., Mixon, R.L., Hobbie, E.A. & Hibbett, D.S. (2011) Conservation of biotrophy in Hygrophoraceae inferred from combined stable isotope and phylogenetic analyses. Mycologia 103: 280–290.
https://doi.org/10.3852/10-195
Siller, I., Kutszegi, G., Takács, K., Varga, T., Merényi, Zs., Turcsányi, G., Ódor, P. & Dima, B. (2013) Sixty-one macrofungi species new to Hungary in ?rség National Park. Mycosphere 4: 871–924.
https://doi.org/10.5943/mycosphere/4/5/3
Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organismal Diversity and Evolution 12: 335–337.
https://doi.org/10.1007/s13127-011-0056-0
Simmons, M.P., Ochoterena, H. & Carr, T.G. (2001) Incorporation, relative homoplasy, and effect of gap characters in sequence-based phylogenetic analysis. Systematic Biology 50 (3): 454–462.
https://doi.org/10.1080/106351501300318049
Smith, A.H. & Weber, N.S. (1980) The Mushroom Hunter’s Field Guide. University of Michigan Press, Ann Arbor.
https://doi.org/10.3998/mpub.20246
Sowerby, J. (1798) Coloured Figures of English Fungi. 2. Printed by J. Davis, London, pp. 121–240.
Staden, R., Beal, K.F. & Bonfield, J.K. (2000) The Staden package, 1998. Methods in Molecular Biology 132: 115–130.
Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1312–1313.
https://doi.org/10.1093/bioinformatics/btu033
Tingstad, L., Gjerde, I., Dahlberg, A. & Grytnes, J.A. (2017) The influence of spatial scales on Red List composition: Forest species in Fennoscandia. Global Ecology and Conservation 11: 247–297.
https://doi.org/10.1016/j.gecco.2017.07.005
Vasas, G. (1993) Old and new methods used for the preservation of fungi in the Hungarian Natural History Museum. Mikológiai Közlemények, Clusiana 32 (1–2): 33–42.
Vincent, Q.V. (2011) ggbiplot: A ggplot2 based biplot. R package version 0.55. Available from: http://github.com/vqv/ggbiplot (accessed 14 February 2019)
Zajta, E. (2012) Distribution of Hygrophorus species in Hungary. Mikológiai Közlemények, Clusiana 51: 223–240.
White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis, M.A., Gelfand, D.H., Sninsky, J.J. & White, T.J. (Eds.) PCR protocols: a guide to methods and applications. Academic Press, New York, pp. 315–322.
https://doi.org/10.1016/B978-0-12-372180-8.50042-1
Arnolds, E. (1990) Tribus Hygrophoreae. In: Bas, C., Kuyper, Th.W., Noordeloos, M.E. & Vellinga, E.C. (Eds.) Flora Agaricina Neerlandica 2. A. A. Balkema, Rotterdam, pp. 115–133.
Babos, M. (1989) The Agaricales s. l. taxa of Hungary. Mikológiai Közlemények, Clusiana 28: 3–234.
Bataille, F. (1910) Flore monographique des hygrophores. Mémoires de la Société d’émulation du Doubs 8 (4): 132–189.
Becker, G. (1954) Un nouvel Hygrophore: Hygrophorus (Limacium) barbatulus. Schweizerische Zeitschrift für Pilzkunde 32: 91–92.
Bohus, G. (1963) New suggestions for preparing fleshy fungi for the herbarium. Mycologia 55: 128–130.
https://doi.org/10.2307/3756389
Bon, M. (1974) Hygrophores du Centre-Est de la France étudiés du salon du Museum 1971. Bulletin mensuel de la Société linnéenne de Lyon 43: 333–343.
Bon, M. (1977) Clé analytique des Hygrophoraceae (suite). Le genre Hygrophorus Fr. Documents Mycologique 7: 25–53.
Bon, M. (1989) Nouveaux taxons (Hygrophoraceae). Documents Mycologique 75: 55–56.
Bon, M. (1992) Die Großpilzflora von Europa. 1. Hygrophoraceae. Übersetzt und bearbeitet von A. Einhellinger, IHW-Verlag, Eching.
Borchsenius, F. (2009) FastGap 1.2. Department of Bio-sciences, Aarhus University, Denmark. Available from: <a href="http://www.aubot.dk/FastGap_home.htm"><span style="text-decoration: underline;">http://www.aubot.dk/FastGap_home.htm</span></a> (accessed 14 February 2019)
Brock, P.M., Doring, H. & Bidartondo, M.I. (2009) How to know unknown fungi: the role of a herbarium. New Phytologist 181: 719–724.
https://doi.org/10.1111/j.1469-8137.2008.02703.x
Bulliard, J.B.F. (1780) Herbier de la France. 2. Chez l’auteur, Didot, Debure, Paris, 48 pp.
Candusso, M. (1997) Hygrophorus s. l. In: Fungi Europaei 6. Libreria Basso, Alassio.
di Marino, E., Montecchio, L. & Agerer, R. (2008) Hygrophorus penarius L. + Fagus sylvatica L. Descriptions of Ectomycorrhizae 11–12: 77–82.
Fernández-Vicente, J., Hidalgo, F. & Oyarzabal, M. (2015) Adiciones al Parque Natural del Gorbea y aledaños, Araba-Bizkaia, País Vasco. Agaricales, Hygrophoraceae (Basidiomycota). Yesca 27: 31–52.
Fries, E.M. (1836a) Corpus Florarum provincialium suecicae 1. Floram Scanicam, Uppsala.
Fries, E.M. (1836b) Anteckningar öfver de i Sverige växande ätliga Svampar. Palmblad, Sebell, Uppsala.
Fries, E.M. (1838) Epicrisis systematis mycologici seu synopsis Hymenomycetum. Uppsala.
Gardes, M. & Bruns, T.D. (1993) ITS primers with enhanced specificity for basidiomycetes–application to the identification of mycorrhizae and rusts. Molecular Ecology 2 (2):113–118.
https://doi.org/10.1111/j.1365-294X.1993.tb00005.x
Gastwirth, L.J., Gel, R.Y., Hui, W.L.W., Lyubchich, V., Miao, W. & Noguchi, K. (2017) lawstat: Tools for Biostatistics, Public Policy, and Law. R package version 3.1. Available from: https://CRAN.R-project.org/package=lawstat (accessed 14 February 2019)
Gouy, M., Guindon, S. & Gascuel, O. (2010) SeaView version 4: a multiplatform graphical user interface for sequence alignment and phylogenetic tree building. Molecular Biology and Evolution 27: 221–224.
https://doi.org/10.1093/molbev/msp259
Hawksworth, D.L. (2013) The oldest sequenced fungal specimen. Lichenologist 45: 131–132.
https://doi.org/10.1017/S0024282912000710
Heim, R. (1948) Deux Hygrophores nouveaux du Jura. Bulletin de la Société Mycologique de France 63: 127–132.
Hesler, L.R. & Smith, A.H. (1963) North American species of Hygrophorus. University of Tennessee Press, Knoxville.
https://doi.org/10.5962/bhl.title.61976
Holec, J. & Wild, J. (2011) Fungal diversity in sandstone gorges of the Bohemian Switzerland National Park (Czech Republic). Czech Mycology 63 (2): 243–263.
Jacobsson, S. & Larsson, E. (2007) Hygrophorus penarioides, a new species identified using morphology and ITS sequence data. Mycotaxon 99: 337–343.
Kalmár, Z. (1966) The frequency of the higher mushroom species in Hungary. Mikológiai Közlemények, Clusiana 4: 17–26.
Kirk, P.M., Cannon, P.F., Minter, D.W. & Stalpers, J.A. (2008) Ainsworth & Bisby’s dictionary of the fungi. 10th edition. CABI International, Wallingford.
https://doi.org/10.1079/9780851998268.0000
Konrad, P. & Maublanc, A. (1937) Icones selectae fungorum, vol 6., Paul Lechevalier, Paris.
Kovalenko, A. (1999) The arctic-subarctic and alpine-subalpine component of the Hygrophoraceae in Russia. Kew Bulletin 54: 695–704.
https://doi.org/10.2307/4110865
Kovalenko, A. (2012) Hygrophorus Fr. In: Knudsen, H. & Vesterholt, J. (Eds.) Funga Nordica. Agaricoid, boletoid, clavarioid, cyphelloid and gastroid genera. Nordsvamp, Copenhagen, pp. 282–293.
Krieglsteiner, G.J. (Ed.) (2000) Die Großpilze Baden-Württembergs. Band 1. Ulmer Verlag, Stuttgart.
Krieglsteiner, G.J. (Ed.) (2001) Die Großpilze Baden-Württembergs. Band 3. Ulmer Verlag, Pliezhausen.
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33 (7): 1870–1874.
https://doi.org/10.1093/molbev/msw054
Larsson, E. & Jacobsson, S. (2004) Controversy over Hygrophorus cossus settled using ITS sequence data from 200 year-old type material. Mycological Research 108: 781–786.
https://doi.org/10.1017/S0953756204000310
Lodge, D.L., Padamsee, M., Matheny, P.B., Aime, M.C., Cantrell, S.A., Boertmann, D., Kovalenko, A., Vizzini, A., Dentinger, B., Kirk, P., Ainsworth, A.M., Moncalvo, J.-M., Vilgalys, R., Larsson, E., Lücking, R., Griffith, G.W., Smith, M.E., Norvell, L.L., Desjardin, D.E., Redhead, S.A., Ovrebo, C.L., Lickey, E.B., Ercole, E., Hughes, K.W., Courtecuisse, R., Young, A., Binder, M., Minnis, A.M., Lindner, D.L., Ortiz-Santana, B., Haight, J., Læssøe, T., Baroni, T.J., Geml, J. & Hattori, T. (2014) Molecular phylogeny, morphology, pigment chemistry and ecology in Hygrophoraceae (Agaricales). Fungal Diversity 64: 1–99.
https://doi.org/10.1007/s13225-013-0259-0
Lowy, B. (1958) On preparing fleshy fungi for the herbarium. Mycologia 50: 442–444.
https://doi.org/10.2307/3756081
Matheny, P.B., Curtis, J.M., Hofstetter, V., Aime, M.C., Moncalvo, J.M., Ge, Z.W., Slot, J.C., Ammirati, J.F., Baroni, T.J., Bougher, N.L., Hughes, K.W., Lodge, D.J., Kerrigan, R.W., Seidl, M.T., Aanen, D.K., DeNitis, M., Daniele, G.M., Desjardin, D.E., Kropp, B.R., Norvell, L.L., Parker, A., Vellinga, E.C., Vilgalys, R. & Hibbett, D.S. (2006) Major clades of Agaricales: a multilocus phylogenetic overview. Mycologia 98: 982–995.
https://doi.org/10.1080/15572536.2006.11832627
Melis, J.L.L., Mir, G. & Prats, M.C. (2017) Aportación Al Catálogo Micológico De Las Illes Balears. Menorca, Ii. Micobotánica-Jaén Año XII Nº1 Enero-Marzo 2017: 66–99.
Nagy, L.G., Kocsubé, S., Csanádi, Z., Kovács, G.M., Petkovits, T., Vágvölgyi, Cs. & Papp, T. (2012) Re-mind the gap! Insertion-deletion data reveal neglected phylogenetic potential of the nuclear ribosomal internal transcribed spacer (ITS) of fungi. PLoS One 7:e49794.
https://doi.org/10.1371/journal.pone.0049794
Osmundson, T.W., Robert, V.A., Schoch, C.L., Baker, L.J., Smith, A., Robich, G., Mizzan, L. & Garbelotto, M.M. (2013) Filling gaps in biodiversity knowledge for macrofungi: contributions and assessment of an herbarium collection DNA barcode sequencing project. PLoS One 8: e62419.
https://doi.org/10.1371/journal.pone.0062419
Peck, C.H. (1898) New species of fungi. Bulletin of the Torrey Botanical Club 25 (6): 321–328.
https://doi.org/10.2307/2478129
Pecoraro, L., Perini, C., Salerni, E. & Dominicis, V.D. (2007) Contribution to the knowledge of the mycological flora of the Pigelleto Nature Reserve, Mt. Amiata (Italy). Flora Mediterranea 17: 143–163.
R Core Team (2015) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.Available from: https://www.R-project.org/ (accessed 14 February 2019)
Rimóczi, I., Siller, I., Vasas, G., Albert, L., Vetter, J. & Bratek, Z. (1999) Magyarország nagygombáinak javasolt vörös listája. Mikológiai Közlemények, Clusiana 38 (1–3): 107–132.
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574.
https://doi.org/10.1093/bioinformatics/btg180
R Studio Team (2015) RStudio: Integrated Development for R. RStudio Inc., Boston, MA. Available from: http://www.rstudio.com/ (accessed 14 February 2019)
Rücker, T. (1990) Die Pilzflora der Gaisbergwälder bei Salzburg, Österreich. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Österreich 127: 165–183.
Seitzman, B.H., Ouimette, A., Mixon, R.L., Hobbie, E.A. & Hibbett, D.S. (2011) Conservation of biotrophy in Hygrophoraceae inferred from combined stable isotope and phylogenetic analyses. Mycologia 103: 280–290.
https://doi.org/10.3852/10-195
Siller, I., Kutszegi, G., Takács, K., Varga, T., Merényi, Zs., Turcsányi, G., Ódor, P. & Dima, B. (2013) Sixty-one macrofungi species new to Hungary in ?rség National Park. Mycosphere 4: 871–924.
https://doi.org/10.5943/mycosphere/4/5/3
Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organismal Diversity and Evolution 12: 335–337.
https://doi.org/10.1007/s13127-011-0056-0
Simmons, M.P., Ochoterena, H. & Carr, T.G. (2001) Incorporation, relative homoplasy, and effect of gap characters in sequence-based phylogenetic analysis. Systematic Biology 50 (3): 454–462.
https://doi.org/10.1080/106351501300318049
Smith, A.H. & Weber, N.S. (1980) The Mushroom Hunter’s Field Guide. University of Michigan Press, Ann Arbor.
https://doi.org/10.3998/mpub.20246
Sowerby, J. (1798) Coloured Figures of English Fungi. 2. Printed by J. Davis, London, pp. 121–240.
Staden, R., Beal, K.F. & Bonfield, J.K. (2000) The Staden package, 1998. Methods in Molecular Biology 132: 115–130.
Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1312–1313.
https://doi.org/10.1093/bioinformatics/btu033
Tingstad, L., Gjerde, I., Dahlberg, A. & Grytnes, J.A. (2017) The influence of spatial scales on Red List composition: Forest species in Fennoscandia. Global Ecology and Conservation 11: 247–297.
https://doi.org/10.1016/j.gecco.2017.07.005
Vasas, G. (1993) Old and new methods used for the preservation of fungi in the Hungarian Natural History Museum. Mikológiai Közlemények, Clusiana 32 (1–2): 33–42.
Vincent, Q.V. (2011) ggbiplot: A ggplot2 based biplot. R package version 0.55. Available from: http://github.com/vqv/ggbiplot (accessed 14 February 2019)
Zajta, E. (2012) Distribution of Hygrophorus species in Hungary. Mikológiai Közlemények, Clusiana 51: 223–240.
White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis, M.A., Gelfand, D.H., Sninsky, J.J. & White, T.J. (Eds.) PCR protocols: a guide to methods and applications. Academic Press, New York, pp. 315–322.
https://doi.org/10.1016/B978-0-12-372180-8.50042-1