Journal des J.E.C. No. 14, p. 7 – 26, 2012 Cortinarius species in acidophilous-eutrophic (but not calciphilous) oak forests of S Norway and Hungary Tor Erik Brandrud *, BálinT dima ** & GEErT SchmidT-STohn *** 1 2 3 received on 26.04.2012 Summary The characteristic, rather species-poor Cortinarius funga of dry, acidophilous to rich but not calcareous oak forests in S Norway and Hungary is compared. Approximately 20 Cortinarius species can be regarded as typical for these forest types in Norway/Hungary, including the following more or less habitat-speciic taxa of non-calcareous Quercus forests: C. cagei, C. emollitus, C. orellanus, C. sabuletorum ss. Saar and C. subcompar. Some further non-calciphilous taxa have a slightly wider host range, and can be regarded as typical Quercus-Carpinus-Fagus species (e.g. C. subporphyropus, C. subpurpurascens) or Quercus-Picea species (C. purpurascens). The Hungarian oak forests differ in the occurrence of a few southern species not found in Norway: C. scaurotraganoides, C. xanthochlorus and C. xanthophyllus. The Norwegian oak forest element includes a few taxa only found in Fagus forests in Hungary: C. barbatus, C. humicola, C. tofaceus. Descriptions are provided for the following rare/little known taxa: C. subpurpurascens, C. subporphyropus, C. sabuletorum ss. Saar, C. xanthochlorus, C. subcompar, C. scaurotraganoides and C. emollitus. C. subcompar is a character species of acidophilous oak forests in Hungary, but is almost unknown elsewhere in Europe, with just a few records from Spain, (probably) France, one from Germany and one from Norway. C. sabuletorum ss. Saar is another, apparently oak forest specialist with very few documented records in Europe. réSumé Les auteurs comparent la fonge du genre Cortinarius caractéristique, plutôt pauvre en espèces, des chênaies sèches, acides à plus riches mais non calciphiles du sud de la Norvège et de Hongrie. Il y a environ 20 espèces de Cortinarius qu’on peut considérer comme typiques de ces forêts en Norvège et en Hongrie, y compris les taxons suivants à habitat plus ou moins spéciique des chênaies non calciphiles : C. cagei, C. emollitus, C. orellanus, C. sabuletorum ss. Saar et C. subcompar. Quelques autres taxons non calciphiles ont une gamme d’hôtes plus large et peuvent être considérés comme des espèces typiques des associations Quercus-Carpinus-Fagus (p. ex. C. subporphyropus, C. subpurpurascens) ou Quercus-Picea (p. ex. Cortinarius purpurascens). Les chênaies hongroises se distinguent par la présence de quelques espèces méridionales qu’on ne trouve pas en Norvège, à savoir C. scaurotraganoides, C. xanthochlorus et C. xanthophyllus. En revanche, on trouve dans les chênaies norvégiennes quelques espèces qui en Hongrie ne poussent que dans les hêtraies : C. barbatus, C. humicola, C. tofaceus. Les auteurs donnent des descriptions * Tor Erik Brandrud, Norwegian Institute for Nature Research (NINA), Gaustadalléen 21, 0349 Oslo, Norway; tor.brandrud@nina.no ** Bálint Dima, Department of Nature Conservation and Landscape Ecology, Faculty of Agricultural and Environmental Sciences, Szent István University, Páter Károly u. 1, H-2103 Gödöllő, Hungary; cortinarius1@gmail.com *** Geert Schmidt-Stohn, Burgstr. 25, D-29553 Bienenbüttel, Germany; geert.schmidt-stohn@t-online.de 7 des espèces suivantes, rares et peu connues : C. subpurpurascens, C. subporphyropus, C. sabuletorum ss. Saar, C. xanthochlorus, C. subcompar, C. scaurotraganoides et C. emollitus. C. subcompar est une espèce caractéristique des chênaies acides en Hongrie, mais à l’exception de quelques mentions en Espagne, probablement aussi en France, et d’une seule en Allemagne et en Norvège, elle est presque inconnue dans le reste de l’Europe. C. sabuletorum ss. Saar est une autre espèce manifestement spécialisée dans les chênaies, très peu mentionnée jusqu’ici en Europe. ZuSammenfaSSung Die charakteristische, recht artenarme Cortinarius-Funga trockener, saurer bis reicherer, aber nicht kalkliebender Eichenwälder Südnorwegens und Ungarns wird verglichen. Annähernd 20 Cortinarius-Arten kann man als typisch für diese Wälder in Norwegen und Ungarn betrachten, einschließlich der folgenden mehr oder weniger standort-speziischen Arten nicht kalkliebender Eichenwälder: C. cagei, C. emollitus, C. orellanus, C. sabuletorum ss. Saar und C. subcompar. Einige weitere nicht kalkliebende Taxa haben ein etwas weiteres Wirtsspektrum und können als typische Quercus-Carpinus-Fagus-Arten (z.B. C. subporphyropus, C. subpurpurascens) oder Quercus-Picea-Arten (z.B. Cortinarius purpurascens) betrachtet werden. Die ungarischen Eichenwälder unterscheiden sich durch das Vorkommen einiger südlicher Arten, die man nicht in Norwegen indet, nämlich C. scaurotraganoides, C. xanthochlorus und C. xanthophyllus. In den norwegischen Eichenwäldern wiederum kommen einige wenige Arten vor, die man in Ungarn nur in Fagus-Wäldern indet: C. barbatus, C. humicola, C. tofaceus. Beschreibungen werden für die folgenden seltenen, wenig bekannten Arten gegeben: C. subpurpurascens, C. subporphyropus, C. sabuletorum ss. Saar, C. xanthochlorus, C. subcompar, C. scaurotraganoides und C. emollitus. C. subcompar ist eine Charakterart saurer Eichenwälder in Ungarn, ist aber mit Ausnahme einiger Nachweise aus Spanien, wahrscheinlich aus Frankreich und je einmal aus Deutschland und Norwegen im übrigen Europa fast unbekannt. C. sabuletorum ss. Saar ist eine andere, offensichtlich auf Eichenwälder spezialisierte Art mit bisher wenigen Nachweisen in Europa. riaSSunto Vengono confrontate specie rare del genere Cortinarius che crescono in boschi di quercia asciutti, da acidi a più arricchiti, ma non calcarei della Norvegia meridionale e dell’Ungheria. Per questi boschi della Norvegia e dell’Ungheria, circa 20 specie del genere Cortinarius sono caratteristiche, comprese le seguenti specie che sono più o meno legate a particolari luoghi di ritrovamento in boschi di quercia di terreni non calcarei: C. cagei, C. emollitus, C. orellanus, C. sabuletorum ss. Saar e C. subcompar. Altri taxa di terreni non calcarei sono micorrizici anche di altri alberi e sono specie tipiche di Quercus-Carpinus-Fagus (per esempio, C. subporphyropus, C. subpurpurascens) o Quercus-Picea-Arten (ed esempio, Cortinarius purpurascens). I boschi di Quercus ungheresi si distinguono per la presenza anche di alcune specie meridionali, che mancano in Norvegia. Queste sono: C. scaurotraganoides, C. xanthochlorus e C. xanthophyllus. Invece, nei boschi di Quercus norvegesi troviamo alcune poche specie che in Ungheria crescono nei boschi di Fagus: C. barbatus, C. humicola, C. tofaceus. Sono descritte anche alcune specie rare e meno note, quali C. subpurpurascens, C. subporphyropus, C. sabuletorum ss. Saar, C. xanthochlorus, C. subcompar, C. scaurotraganoides e C. emollitus. C. subcompar è una specie rara, tipica dei boschi acidi di Quercus in Ungheria, ma eccetto alcuni ritrovamenti segnalati in Spagna, probabilmente anche in Francia e un unico ritrovamento in Germania e in Norvegia, è una specie quasi sconosciuta nel resto d’Europa. Cortinarius sabuletorum ss. Saar è un’ulteriore specie verosimilmente legata ai boschi di Quercus, ma inora sono segnalati solo pochi ritrovamenti in Europa. 8 Introduction Calcareous deciduous forests house a very diverse funga of Cortinarius species. Many taxa especially of subgenus Phlegmacium are calciphilous and many are more or less restricted to calcareous forests dominated by Quercus, Carpinus, Fagus, Tilia and Corylus (Brandrud & BEndikSEn 2001, Brandrud & SchmidT-STohn 2011, JEppESEn & FrøSlEv 2011). Deciduous forests on less or non-calcareous soils are much less species rich in cortinarii, and their species composition differs considerably (cf. KnudSEn & VESTErholT 2008). However, the Cortinarius funga of the non-calcareous forests are less focused and less documented. In the present paper, we compare the acidophilous to eutrophic oak-dominated forests on noncalcareous ground in S Norway and Hungary. Both regions, although widely separated, have welldeveloped oak forests as a major component on thermophilic–xerophilic sites with intermediate to acid bedrocks, and in both regions these forests have been extensively mapped for Cortinarius species. Material and Methods Data from the Norwegian and Hungarian museum herbaria (herb. O, herb. BP; including own material) has been compiled, with focus on data on habitat preference. Furthermore, the presented species have been studied from various oak forests of the regions, including documentations with photographs and descriptions of macro- and micromorphology. Lists of investigated material from Norway and Hungary are included, with the present authors abbreviated TEB, DB and SSt. Some of the collections have also been DNA (ITS) analysed, following standard methods (FrøSlEv et al. 2005, cf. also SchmidT-STohn & OErTEl 2009). Eutrophic–acidophilous oak forests in S Norway and Hungary S Norway: Xerophilous-thermophilous Quercus robur-Q. petraea forests are found along the coast of southernmost Norway, with a concentration in the counties of Aust-Agder and Vest- Photo: Tor Erik Brandrud Fig. 1 — Mesotrophic, low-herb oak forest in S Norway. Ytre Lauvrak, Froland, Aust-Agder County 9 Agder. These are usually open, dry, shallow-soil forest types (Fig. 1), with co-dominance of Corylus avellana and Tilia cordata (in rich types) as well as Populus tremula. Carpinus betulus is often co-dominant in nemoral European oak forests, but does not occur as far north as Norway. These dry oak forests range from edaphic rich (but not calcareous) low herb types with species such as Hepatica nobilis, Lathyrus niger and (abundant) Convallaria majalis to strictly acidophilous ones with Deschampsia lexuosa, Melampyrum pratense and Vaccinium myrtillus. Most stands occur on siliceous, but somewhat richer bedrocks (amphibolites, larvikites, basalts), whereas the most acid bedrock ridges are occupied by pine forests. These thermophilous oak forests include a number of habitat-speciic, southern fungi in Norway, such as Hydnellum and Ramaria species (see e.g. Brandrud 2007, Brandrud et al. 2012). Floristically, the richest Quercus-Tilia forest types on more siliceous bedrocks («amphibolite oak-lime forests») resemble the calcareous Tilia forests of the Oslofjord area, but their fungal elements area quite different (cf. lists of calcareous Tilia forest species in Brandrud et al. 2011). Hungary: Four main association types of the acidophilous oak forests exist in Hungary (Genisto pilosae-Quercetum petraeae, Deschampsio lexuosae-Quercetum sessililorae, CastaneoQuercetum and Luzulo-Carpinetum). They are developed mainly on siliceous bedrocks, e.g. amphibolite, rhyolite, dacite, andesite, gneiss, mica, sandstone, etc., usually at the top of steep slopes (25–40°) (Fig. 2). All of them are characterized by Quercus petraea mixed with e.g. Car- Photo: Bálint Dima Fig. 2 — Acidophilous oak forest in Hungary on sandstone. Mt Ezüst-hegy, Pilis Mts, Budakalász, Pest County pinus betulus, Fagus sylvatica at higher altitudes/N exposition, and with Quercus robur, Castanea sativa, Pinus sylvestris in W and SW Hungary. Quercus pubescens might be co-occurring on more thermophilic sites, and Betula pendula and Populus tremula in younger stands. The poor vegetation is characterized by e.g. Deschampsia lexuosa, Genista pilosa, Hieracium spp., Luzula luzuloides, Melampyrum pratense, Vaccinium myrtillus. The moss layer is often well-developed sometimes with almost 100% coverage, with Dicranum spp., Polytrichum spp. and Leucobryum glaucum. The funga of these forest types are dealt with in the classic mycoenological study of BohuS and BaBoS (1967) and BohuS (1984), focusing the fungal assemblages along the gradient of soil pH from calciphilous to acidophilous frondose forests. 10 Cortinarius species with preference for eutrophic-acidophilous oaks forests S Norway: Altogether 15 Cortinarius species have the majority of Norwegian records in dry, eutrophic-mesotrophic(-acidophilous) oak forests (Tab. 1). This list includes e.g. three taxa in each of the (sub)sect. Purpurascentes and sect. Phlegmacioides of subgen. Phlegmacium, and ive taxa of subgen. Telamonia. A few species are rather frequent (C. largus/lividoviolaceus, C. orellanus, C. torvus), but most are fairly rare, with <15 known occurrences under oak. Many of these preferent oak forest species also have a number of localities in the steep scree forests of Tilia(-Corylus) type or Corylus (-Betula-Pinus) type along the fjords of W Norway, some also in SE Norwegian Fagus forests. However, this fungal community normally lacks in the calciphilous Tilia-Corylus forests in SE Norway (Tab. 1). Tab. 1. Cortinarius species with preference for mesotrophic to rich (but not calcareous) oak forests in S Norway (I = rich/mesotrophic Quercus (-Tilia) forests; II = rich Tilia scree forests; III = rich Corylus scree forests; IV = rich/mesotrophic Fagus forests; V = calcareous Tilia (-Quercus) forests; VI = calcareous Corylus forests; VII = other forest types). The number of known occurrences/localities in various thermophilous deciduous forest types is indicated. Species more or less exclusively associated with rich/mesotrophic oak forests in Norway indicated in bold. Cortinarius subgenus Phlegmacium C. balteatocumatilis C. largus (C. lividoviolaceus ss. Saar) C. purpurascens ss. str. C. sabuletorum ss. Saar* C. subporphyropus C. subpurpurascens [C. talus] [C. variecolor] I II III IV V VI VII TOTAL 16 2 6 2 3 3 7 40 45 20 26 9 3 3 18 5 6 15 15 12 4 9 3 11 5 3 1 4 10 2 1 11 3 3 106 18 3 2 27 130 1 2 3 18 1 50 5 10 47 60 150 subgenus Telamonia C. acetosus (C. rigens ss. Lange) C. cagei C. subcompar C. torvus 11 4 1 24 1 2 4 3 3 8 55 1 4 17 7 1 50 subgenus Cortinarius C. humicola C. tofaceus C. orellanus 3 10 60 1 1 subgenus Myxacium C. barbatus coll. 7 C. emollitus 9 *=C. latobalteatus in kålåS et al. (2010) 6 1 2 2 2 16 13 Altogether 9 species are exclusively or almost exclusively associated with rich to acidophilous oak forests, including three taxa often placed in section Orellani of subgen. Cortinarius (C. humicola, C. orellanus, C. tofaceus). Most of the species in Tab. 1 are found in the rich oak forest types, some in mesotrophic/semi-rich ones, and only few species such as C. emollitus occur in the strictly oligotrophic/acidophilous types in Norway. Cortinarius talus and C. variecolor, being most frequent in Betula pubescens and Picea abies forests, resp., are also included in Tab. 1, since their southern, thermophilous deciduous forest populations have a preference for eutrophic-acidophilous oak forests. 11 Hungary: A total of 15 Cortinarius species are typical in acidophilous oak forest type in Hungary (Tab. 2), from which 12 are also found in similar habitats of Norway. These cortinarii are more or less rare (except C. torvus, C. cagei) but widespread. Altogether 9 species (C. balteatocumatilis, C. purpurascens, C. sabuletorum ss. Saar, C. xanthochlorus from subgen. Phlegmacium; C. cagei, C. scaurotraganoides, C. subcompar from subgen Telamonia; C. orellanus from subgen. Cortinarius; and C. emollitus from subgen. Myxacium) are almost exclusively associated with oaks on acid to strongly acid soil, and can be regarded as character species of these oak forests. However, all of the species included in Tab. 2 can also occur occasionally in acidophilous Fagus forests, or in Picea plantations in case of C. purpurascens. Cortinarius largus, C. subporphyropus and C. talus are fairly rare with Quercus, but more common with other hosts e.g. Fagus or Betula. The occurrence of C. variecolor and C. acetosus (C. rigens ss. Lange) is uncertain due to the confusion of the related species. Tab. 2. Cortinarius species with preference for acidophilous-eutrophic oak forests in S Norway and Hungary. xx = more or less exclusively associated with acidophilous-eutrophic oak forests. X(X) = most frequent habitat. X = present. Species subgenus Phlegmacium C. balteatocumatilis C. largus C. purpurascens ss. str. C. sabuletorum ss. Saar C. subporphyropus C. subpurpurascens C. talus C. variecolor C. xanthochlorus C. xanthophyllus S Norway Hungary X(X) X(X) X(X) xx X(X) X(X) (X) (X) – – xx X xx xx X X(X) X ? xx X xx xx xx* X(X) ? xx xx xx X xx xx xx – – xx X(X) xx 15 xx 15 subgenus Telamonia C. acetosus (C. rigens ss. Lange) C. cagei C. scaurotraganoides C. subcompar C. torvus subgenus Cortinarius C. humicola C. tofaceus C. orellanus subgenus Myxacium C. barbatus coll. C. emollitus TOTAL (20 species NO+HU) *hitherto recorded only once in Norway. Comparison of the cortinarii of oak forests in S Norway versus Hungary The acidophilous-eutrophic oak forest element is remarkably similar in S Norway and Hungary; at least 12 of the species are common for these kinds oak forests in the two areas, and almost all of the most specialized oak forest species in S Norway occurs in similar oak forests in Hungary. 12 At least the following taxa can be regarded as specialized non-calcareous Quercus forest species in both countries: C. cagei, C. emollitus, C. orellanus, C. sabuletorum ss. Saar and C. subcompar. The Hungarian oak forests differ, however, in the occurrence of some southern species not found in Norway; C. scaurotraganoides, C. xanthochlorus and C. xanthophyllus. Among the taxa on the Norwegian list, C. humicola, C. tofaceus and C. barbatus cannot be found in Hungarian oak forests, but these occur in other acidophilous-mesotrophic forest types in Hungary. Some of the species treated here, can be regarded as typical Quercus-Carpinus-Fagus species on noncalcareous ground (e.g. C. subporphyropus, C. subpurpurascens). The high similarity in the eutrophic-acidophilous oak forest element in S Norway and Hungary indicates that this fungal element is widespread, and with fairly similar habitat requirements in different parts of Europe, ranging from northwestern, suboceanic areas to central, continental districts. The same element is apparently found also in SW Germany; in park-like oak stands on loamy/sandy acid soils, with inds of C. sabuletorum, C. lividoviolaceus, C. balteatocumatilis, C. subpurpurascens, C. orellanus, C. hinnuleus, C. torvus, C. subcompar and C. emollitus (G. Saar, pers. comm.). Cortinarius subcompar, which stands out as one of the more habitat-speciic taxa of acidophilous oak forests, is recorded from this habitat also in the Thüringen district of Germany (pers. obs. SSt; Kyffhäuser/Rothenburg; acidophilous oak forest on sandstone with Leucobryum glaucum; together with C. bolaris), and apparently also occur in similar habitat in oceanic areas of Spain (CadiñanoS AGuirrE 2006; sub nom. C. moserianus, C. pseudophlegma). However, some differences in habitat between Hungary and S Norway can be seen. There is a tendency that these species occur in more acidophilic oak forest types in Hungary than in Norway. For instance, the poisonous and well-known C. orellanus occurs rather strictly in acidophilous oak forests in Hungary. Otherwise in C Europe it is partly indicated as a character species for acido-thermophilous habitats (GmindEr & KriEGlSTEinEr 2010, DähnckE & DähnckE 1979) and partly indicated from basic as well as acidic soils (BrEiTEnBach & Kränzlin 2000), whereas it in Norway almost always grows in the richer (but not calcareous) low herb oak forests, often together with Hepatica nobilis. This regional difference is even more pronounced in the case of the yellow/greenish Phlegmacium species C. xanthophyllus and C. xanthochlorus. BohuS (1970, 1984) documented that these were growing in more or less acidophilic forests with low pH topsoils in Hungary. In the northernmost outposts for these species in SE Sweden, on the contrary, these are occurring on strictly calcareous ground. Similar regional patterns are seen also with some other species in related habitats. For instance, Cortinarius splendens occurs mainly on siliceous ground under Quercus suber in Mediterranean areas (pers. obs. TEB), whereas the species in NE Germany occurs in calcareous beech forests (Brandrud & SchmidT-STohn 2011) and in SE Norway in calcareous Tilia forests (Brandrud et al. 2011). Descriptions Cortinarius subpurpurascens (Batsch) Fries Fig. 3, 4 Cap 5–10(–12) cm, broadly umbonate, near margin usually with hygrophanous ring zone, spots or radially veins, fulvous brown to rather pale ochraceous brown, incurved margin greyish white to rarely bluish tinged. Gills pale greyish violet, turning somewhat vinaceous when bruised. Stem 6–10 × 1.2–2.5 cm, bulb up to 3 cm wide, bulb clavate to faintly marginate, often pointedtapering, pale greyish blue-bluish grey, pale vinaceous-lilac on bruising. Flesh pale greyish violet in stem, whitish in cap, when young lilac on bruising. Smell distinctly honey-like. Chem. reactions: lesh purplish with lugol. 13 Spores 8.5–9.5(–10) × 5–6 µm, ellipsoid, very strongly and medium coarsely verrucose. Cap cuticle duplex, subcutis/hypoderm with cellular structure and yellow thick wall (parietal) pigment. DNA (ITS) sequence: High similarity to C. purpurascens. No infraspeciic variation observed, but according to G. Saar (pers. comm.), another genotype exists, found from calcareous area. Photo: Kristin H. Brandrud Fig. 3 — Cortinarius subpurpurascens (Norway), TEB 775-11 Photo: Geert Schmidt-Stohn Fig. 4 — Cortinarius subpurpurascens (TEB 775-11) spores 2000 : 1, focus-stacking from 9 single photos 14 Habitat and distribution: Eutrophic to acidophilous Quercus, Carpinus, Fagus, Tilia and Corylus forests, very rarely on calcareous ground. Apparently distributed in most parts of Europe; from NW Norway-C Sweden south to the Mediterranean area. Distribution in some areas uncertain due to confusion with the related C. purpurascens. Investigated material from NO and HU: NORWAY: Østfold; Fredrikstad, TEB 1980. Oppland; Lunner, TEB 442-08 (Corylus); TEB 775-11 (Corylus). Akershus; Frogn, TEB 9-89, 33-94 (Tilia-Quercus); Asker, TEB 77-92 (Tilia-Quercus); Bærum TEB 498-04 (Corylus). Buskerud; Øvre Eiker, TEB 195-85 (Corylus). Vestfold; Larvik, TEB 382-08 (Quercus-Tilia); TEB 30210 (Quercus-Tilia). Aust-Agder; Tvedestrand, TEB 48-01 (Quercus-Tilia). Rogaland; Suldal, TEB 175-00 (Tilia-Corylus). Hordaland; Bergen, TEB 310-80, 427-80 (Fagus); Ulvik, TEB 361-05, 443-05 (Tilia). HUNGARY: Budapest, DB 2847 (Quercus). Nógrád; Diósjenő, DB 4194 (Fagus, Carpinus). Vas; Szalafő, DB 4225 (Quercus, Carpinus, Fagus). Szabolcs-Szatmár-Bereg; Vámosatya, DB 3751 (Quercus). Comments: C. subpurpurascens is distinguished from C. purpurascens by the more clavate, hardly-marginate bulb and the often paler, more ochraceous brown pileus. Generally, the violaceous colours are also paler (and usually lacking on cap) on C. subpurpurascens. The spores are furthermore slightly larger. The taxon has formerly often been named C. purpurascens var. largusoides. C. purpurascens have a wider ecology, occurring in deciduous and in coniferous forests, including young Picea plantations. In oak forests, however, these taxa may be co-occurring. C. subpurpurascens in Norway normally occur in rich low-herb Quercus(-Tilia-Fagus) forests. However, it is a «negative indicator» of the calciphilous Cortinarius element; when you ind this species, you normally do not ind the calciphilous ones. In Hungary it is typical in acidophilous or subacidophilous frondose forests, associated mainly with Quercus, but also with Fagus and Carpinus. Cortinarius subporphyropus Pilát Fig. 5, 6 Cap 2–4 cm, umbonate, innately ibrillose, initially pale brownish grey-bluish grey, later silvery white to more ochraceous brown at centre. Gills greyish violet, turning darker lilac when bruised. Stem 2.5–5 × 0.3–0.6 cm, bulb up to 1 cm wide, pale greyish violet, pale lilac on bruising. Flesh bluish in stem, bulb and cap lesh almost whitish; lilac on bruising. Smell distinctly honey-like. Chem. reactions: purplish with lugol. Spores (9–)10–11.5(–12) × 5.5–7 µm, ellipsoid, strongly and densely verrucose. Cap cuticle duplex, subcutis/hypoderm with cellular structure and pale yellow thick wall (parietal) pigment. DNA (ITS) sequence: No infraspeciic variation observed. High afinity to C. porphyropus. Habitat and distribution: In mesotrophic to oligotrophic/acidophilous Quercus-Carpinus, Tilia and Fagus forests, sometimes also in Picea plantations. In Hungary also with the same hosts but twice found under Betula and/or Populus tremula, where Quercus was also near! Widespread, from NW Norway to Hungary and France (pers. obs. TEB, DB). Apparently rare everywhere. Investigated material from NO and HU: NORWAY: Oppland; Lunner, TEB 441-08 (Corylus). Akershus; Bærum, TEB 142-01 (Tilia-Quercus). Vestfold; Horten, TEB 407-78 (Fagus); Tønsberg, TEB 257-80, P. Marstad 1820911 (Fagus). Telemark; Kragerø, TEB 239-87 (Quercus). Aust-Agder; Evje & Hornnes, TEB 298-00, 299-00 (Tilia); Tvedestrand, TEB 216-01 (Quercus). Hordaland; Ulvik, TEB 249-07 (cf.) (Tilia, Corylus). HUNGARY: Somogy; Böhönye, DB 1386 (Quercus, Carpinus, Tilia). Vas; Szalafő, DB 2134 (Betula, Populus tremula, Quercus). Pest; Pilisszentlászló, DB 3360 (Fagus). 15 Photo: Bálint Dima Fig. 5 — Cortinarius subporphyropus (Hungary), DB 2134 Photo: Geert Schmidt-Stohn Fig. 6 — Cortinarius subporphyropus (SSt 07-212) spores 2000:1, focus-stacking from 6 single photos 16 Comments: Cortinarius subporphyropus is the smallest Phlegmacium species in Europe. The species is distinguished from C. porphyropus by the smaller fruitbodies and the larger spores (C. porphyropus spores: 8–9.5 × 5–6 µm). The habitat is also different; C. porphyropus is a strict Betula associate, mainly occurring in N Europe subalpine Betula forests, but also with Betula in C Europe, whereas C. subporphyropus mainly occur in Quercus and Fagus forests, sometimes also in Picea plantations, possibly also with Betula. The spore morphology of C. subporphyropus shows a high, but apparently not taxonomic relevant variation. In S Norway, comparatively small spore are found, 9–10.5 × 5.5–6.5 µm, whereas in Hungary spores (9.5–)10–11.5(–12) × 5.5– 7 µm are found. Cortinarius mendax Bidaud, Mahiques & Reumaux is macroscopically quite similar to our species, but is described with even smaller spores, of the size of C. porphyropus spores (8–10 × 5–6 µm; Bidaud 2011). Cortinarius sabuletorum Redeuilh & Reumaux ss. Saar Fig. 7, 8 Cap 4.5–8(–10) cm, convex, viscid-glutinous when young, often radially innately ibrillose, initially (pale) greyish blue, at least towards margin, sometimes completely whitish, but soon discolouring to pale greyish brown, centre becoming more fulvous brown, sometimes apparently almost whitish towards margin. Gills initially greyish white. Stem 4–8 × 1–2 cm, bulb up to 2.5 cm wide, cylindrical-clavate to almost marginate bulbous, greyish white, somewhat (red) brownish spotted with age; when young girdled or peronate of lilac blue veil. Flesh whitish, sometimes with a faint bluish tinge in cap, slightly or distinctly browning or brown spotted when bruised. Smell somewhat to strongly earth-like (variecolor-smell). Chem. reactions: KOH in the lesh strong yellow-golden yellow, after half an hour turning pinkish brown. Photo: Bálint Dima Fig. 7 — Cortinarius sabuletorum ss. Saar (Hungary), DB 4324 17 Spores 9.5–11.5 × 5.5–6.5 µm, amygdaloid to faintly citriform, medium strongly and rather densely verrucose. Cap cuticle more or less simplex, surface hyphae distinctly gelatinous, erectentangled. Photo: Geert Schmidt-Stohn Fig. 8 — Cortinarius sabuletorum (SSt 10-161) spores 2000:1, focus-stacking from 6 single photos. The ornamentation is very lat, therefore the ornaments are hardly visible at the outline of the spores DNA (ITS) sequence: No internal (infraspeciic) variation found. Differs by >30 nucleotides to related species. Nearest neighbour species not clear. Habitat and distribution: In rich-mesophilous-acidophilous (non-calcareous) QuercusCarpinus forests. Hitherto with certainty known from S Norway, Hungary, Germany (Saar 2010) and France (Jura; pers. obs.; cf. also Bidaud et al. 1995, 1996, sub nom. C. sabuletorum, C. clarobaltoides). Investigated material from NO and HU: NORWAY: Vestfold; Larvik, TEB 455-08, 46208 (Quercus-Tilia). Telemark; Kragerø, TEB 529-09, 911-11, 912-11 (Quercus). Aust-Agder; Arendal, TEB 640-11 (Quercus). HUNGARY: Budapest, DB 4324 (Quercus). Heves; Tarnalelesz, DB 1160 (Quercus, Carpinus). Heves; Parádóhuta, leg. L. Albert, AL 10/352 (Quercus). Comments: C. sabuletorum ss. Saar is characterized by its initially pale bluish cap (margin) and veil on stem, and otherwise whitish stem, lesh and gills. These features resembles those of C. balteatocumatilis, but C. sabuletorum ss. Saar is generally much paler, usually smaller and becomes more pronounced brown discoloured when bruised. The present species also resembles C. largus, but this always has a (faint) tinge of bluish in young gills and stem. The species has been quite unknown until recently, but was depicted and described in detail by Saar (2010). 18 The blue pigment is fugacious and seems mainly restricted to the veil. Saar (2010) described C. sabuletorum with cap «jung weisslich, graulich-lila, lilabräunlich», and one picture shows young specimens with white cap margin. In our experience, the species normally has a pale lilacbluish or pale greyish blue cap when young and quite fresh, as a «diluted C. balteatocumatilis», but quite white young caps have been observed occasionally also in Hungary (see Fig. 7). The original description of C. sabuletorum (Bidaud et al. 1995) its our species in many respects, but might also represent pale forms of C. largus or related species without bluish tinges, and the type material should be further investigated (including DNA). Cortinarius clarobaltoides Rob. Henry ss. Bidaud et al. (Bidaud et al. 1996: Pl. 207), probably also represents our species. The species is typical for eutrophic-acidophilous oak-hornbeam forests in Norway and Hungary, and is reported from the same habitat also in Germany (Saar 2010: «auf nicht kalkhaltigen Böden»). The species are named C. latobalteatus in the Norwegian red list (KålåS et al. 2010). Cortinarius xanthochlorus Rob. Henry Fig. 9 Cap 4–8(–10) cm, convex, with age often slightly innately ibrillose, when young yellowish green or pale olive green, centre more olive brown. Gills yellowish green. Stem 4–6 × 1.2–2.5(–3) cm, with an often acutely marginate bulb (–5 cm), greenish yellow, veil at bulb margin slightly volva-like, greenish yellow. Basal mycelium whitish, sometimes with a lilac tinge. Flesh greenish yellow in stem, whitish in pileus and bulb. Macrochemical reactions: KOH olive brown in context and on surfaces, 40% KOH often faintly reddish on bulb margin. Smell faint, somewhat unpleasant, yeast-like. Photo: Bálint Dima Fig. 9 — Cortinarius xanthochlorus (Hungary), DB 1161 Extractable pigments: lavomannin-6,6’-dimethylether (FDM). Spores 12–14 × (6.5–)7–8 mm (MV = 13.14 × 7.36 µm), Q = 1.79, citriform to amygdaliform, very distinctly and rather densely verrucose. Cap cuticle ± simplex, hyphae (3–)4–8(–10) µm 19 wide, some hyphae zebra-striped encrusted, below with cemented bundles with thick, refractive walls. Also with greenish yellow pigment, becoming ± purple with KOH on exsiccates. DNA (ITS) sequence: Phylogenetically apparently fairly isolated; differing >30 nucleotides to related species. Habitat and distribution: Nemoral and (sub)mediterranean deciduous forests. Associated mainly with Quercus spp., mainly Q. petraea, Q. cerris and Q. pubescens, more rarely with Q. robur, and Fagus sylvatica, probably also with Carpinus betulus, in thermophilous, calciphilous to mesophilopusacidophilous, often dry, grazed, herb-rich, open woodlands or forests of Quercus or Quercus-Carpinus type. Very rare. Most records are from Hungary (e.g. BohuS 1970 as C. olivascentium p.p.; Rimóczi & AlBErT 1992, AlBErT 1996, VaSaS & LocSmándi 2009) and the Appennines, C Italy (PödEr 1982; ConSiGlio 1997 as C. olivascentium and C. pseudofulgens). Otherwise recorded in France (HEnry 1966; EySSarTiEr 2004), Spain (Ballarà 1999, as C. olivascentium), Belgium (VEllinGa 1986), Switzerland, Germany (KriEGlSTEinEr 1991) and a few outpost localities in Sweden (Brandrud et al. 1996). Depicted material from Japan apparently also belong here (leg. & det. K. Maruyama, available on the Internet). Investigated material from HU (and some reference collections): SWEDEN. Öland, Högsrum, Halltorps hage, S. Jacobsson 94057 (Quercus; see Brandrud et al. 1996) (GB). FRANCE. Doubs, R. Henry 508 (holo., hb. Henry). ITALY. Emilia-Romagna, Parma, Borgotaro, Stabielle, TEB 363-81, TEB 387-81 (Quercus; see PödEr 1982). HUNGARY: Zala; Murarátka, leg. G. Bohus & M. Babos, BP 32789 (Quercus, Carpinus) (as C. olivascentium; BohuS 1970). Borsod-AbaújZemplén; Égerszög, leg. G. Vasas, Cs. Locsmándi, G. Bohus, A. Bathó, BP 90978 (Quercus, Carpinus, Fagus) (VaSaS & LocSmándi 2009). Borsod-Abaúj-Zemplén; Háromhuta (Újhuta), AL 95/59 (Quercus) (AlBErT 1996, as C. olivascentium). Heves; Tarnalelesz, DB 1161 (Quercus, Carpinus). Heves; Parádóhuta, DB 1780 (Quercus). Pest; Szentendre, DB 3336 (Quercus). Comments: C. xanthochlorus is characterized by its olive green colours, robust fruitbodies with a sharply marginated bulb, innately ibrillose cap and large spores. The spores are larger than those of any other, resembling frondose forest taxa. The colours resembles those of C. citrinus, and their main pigment lavomannin-6,6’-dimethylether (FDM) is the same, but these two species are genetically fairly distant according to ITS sequences (FrøSlEv et al. 2007). The species seems restricted mainly to southern temperate-supramediterranean, open oak-hornbeam forests. The species is collected form calcareous soils e.g. in the Appennines in C Italy, in W Switzerland, S Germany and S Sweden, whereas in Hungary it occurs in more siliceous soils, almost avoiding the calciphilous forests types. The latter habitat preference is untypical for the calochroid-fulvoid Phlegmacium species. The present concept its well with the type material of C. xanthochlorus, having the characteristic, large spores, and FDM pigment. According to the descriptions in HEnry (1951, 1966), the name C. olivascentium seems to be a synonym, but the spores in the type material are too small. Cortinarius subcompar Bohus Fig. 10, 11 Cap 3–6 cm, convex, often slightly umbonate, silky ibrillose when young, dry but may be slightly sticky when wet, hygrophanous, greyish-bluish to ochraceous brownish, sometimes pale silvery greyish, usually with persistently bluish tinges at margin. Gills crowded, emarginate, pale greyish blue at irst, blush tinges often very weak, and soon fading, then turning pale ochrerusty brown. Stem 2–4 × 0.5–1 cm, with clavate to marginate bulb at the base, dry, whitish ibrillose, with bluish tinges, especially at apex (often contrasting more greyish lamellae); cortina pale greyish-blue. Flesh greyish, bluish-violet, with hygrophanous streaks, becoming ochre at the stem base, taste slightly sour, smell indistinct. Chem. reaction: brownish with KOH. 20 Photo: Bálint Dima Fig. 10 — Cortinarius subcompar (Hungary), DB 3118 Photo: Geert Schmidt-Stohn Fig. 11 — Cortinarius subcompar (SSt 07-190) spores 2000:1, focus-stacking from 6 single photos 21 Spores (6.5–)7–8.5 × 4.2–5 µm, ovoid to (broadly) ellipsoid, strongly and rather densely verrucose, moderately to fairly strongly dextrinoid. DNA (ITS) sequence: Preliminary DNA (ITS) study shows that C. subcompar is related to C. malachius (Fr.) Fr. and C. quarciticus H. Lindstr., both associated with conifers. Habitat and distribution: In acidophilous nemoral and submontane frondose forests, associated with Quercus spp. (rarely with Fagus sylvatica), somewhat thermophilous, main fruiting period is at the end of August or beginning of September. Norwegian ind from rich Quercus-Tilia forest. C. subcompar is a character species in the Hungarian acidophilous Quercus forests, especially at the Vaccinium-Leucobryum type. Rare but widespread in Hungary (from W Hungary to NE Hungary), however, it is almost completely unknown in other European countries. Recently recorded from Germany (Saar, Schmidt-Stohn pers. com.), Norway and from Spain (CadiñanoS AGuirrE 2006, sub nom. C. moserianus and C. pseudophlegma; in acidophilous oak forests), possibly also found in France (HEnry 1981, sub nom. C. pseudophlegma). Investigated material from NO and HU: NORWAY: Aust-Agder, Arendal, leg. I. Kytövuori, TEB 625-11 (Quercus-Tilia). HUNGARY: Pest; Budakeszi, BP 56933 (holotype) (Quercus). Pest; Budakeszi, DB 1775 (Quercus); Győr-Moson-Sopron; Sopron, DB 3118 (Quercus); Heves; Parádóhuta, DB 4124 (Quercus, Fagus). Comments: Cortinarius subcompar is characterized by the stout and phlegmacioid appearance, the silvery greyish-bluish colours, the persistently low umbo on cap, the almost marginated bulb, the small spores and the occurrence on acid soils with frondose trees. Hungary seems to be the chore area for this rare and little known species. In Hungary the species can be easily recognized since no similar species occur in that kind of habitats. The coniferous species, C. quarciticus seems to be closely related according to morphology and DNA studies. In CadiñanoS AGuirrE (2006), the described and depicted material of C. moserianus and C. pseudophlegma, possibly also C. argentatum-silvae represent C. subcompar. Furthermore, the original description of C. pseudophlegma Rob. Henry its also well with C. subcompar (HEnry 1981, p. 272–273.), thus the former seems to be a synonym of the latter. But further molecular studies are needed to clarify this hypothesis. Cortinarius scaurotraganoides Rob. Henry ex Rob. Henry Fig. 12 Cap 5–9 cm, convex, ibrillose-tomentose when young, later becoming naked, radially marbled, may be slightly sticky only when wet; whitish-cream, greyish, pale ochre, sometimes with faint bluish tinges at margin, ochre brown when wet, sometimes slightly hygrophanous with age. Gills broad, crowded, emarginate; ochre brown at irst, inally rusty brown, edge paler, somewhat wavy. Stem 5–8 × 1.5–2 cm, light greyish ochre, with more or less marginated bulb (2.5–4 cm wide), with abundant ibrillose silver grey veil remnants at bulb margin. Flesh yellowish ochre, sometimes with violet tinges at stemapex, saffron to rusty ochre brown at the base; taste sour, smell strongly fruity, like that of C. traganus. Spores 7–8.5(–9) × 4.5–5.5 µm, ellipsoid to somewhat amygdaloid, inely verrucose, moderately dextrinoid. Habitat and distribution: In acidophilous Quercus forests (Vaccinium and Leucobryum type), only exceptionally in Fagus-Carpinus forests, with distinct southern and thermophilous distribution. Associated with different species of Quercus, e.g. Q. petraea, Q. pubescens, Q. suber, and very rarely with Carpinus or Fagus. Known from France (Bidaud et al. 1993), Hungary (LukácS 2002, AlBErT 2011) and Spain (Ballarà et al. 2010). Apparently very rare, but seems to be widespread in S Europe. 22 Investigated material from HU: HUNGARY: Győr-Moson-Sopron; Sopron DB 3117, DB 3703 (both Quercus). Vas; Orfalu, DB 131-2-5-1 (Quercus, Fagus). Pest; Budakeszi, leg. L. Albert, AL 10/325 (Quercus) (AlBErT 2011). Pest, Tahi, leg. L. Albert & DB; AL 10/427 (Quercus). Photo: Bálint Dima Fig. 12 — Cortinarius scaurotraganoides (Hungary), DB 3117 Comments: Cortinarius scaurotraganoides is characterized by the whitish-greyish colours, the saffron or orange tinged lesh at the stem base, the strong and typical smell of C. traganus, and the occurrence in acidophilous frondose forests. It resembles to C. traganus but the latter differs by having distinct bluish colours, slightly larger spores (8–9.5 × 5–6 µm) and different ecology (boreal and montane coniferous forests). C. traganus f. ochraceus is very similar, but it also has somewhat larger (traganus-like) spores and an associated with conifers. Cortinarius niveotraganus Kytöv. Niskanen & Liimat. ad int. is also member of this group, but it occurs in boreal Betula forests and has distinctly larger and strongly verrucose spores. Cortinarius emollitus Fr. Fig. 13 Cap 2–7 cm, convex to umbonate, margin often undulate; viscid-glutinous, innately ibrillose (at least when not too dry), due to ine hygrophanous veins; vividly ochraceous brown to yellow, sometimes with orange tinges, often uniformly coloured, or somewhat more brown at centre, margin initially often pale ochraceous yellow to yellow white. Gills initially greyish white, later greyish brown, pale brownish. Stem 3–8 × 0.5–1.5 cm, bulb up to 2 cm wide, cylindrical-clavate to strongly bulbous, sometimes with a faintly marginate bulb, sometimes tapering below the bulb, slender to (very) robust, whitish, persistent or somewhat ochraceous with age; universal veil (very) thin, viscid, white, restricted to zone near bulb. Flesh whitish, taste bitter in all parts. Smell indistinct. Spores: (6.5–)7–8.5 × 4–5(–5.5) µm ellipsoid to broadly ellipsoid, weakly verrucose. 23 DNA (ITS) sequence: Identical sequence found on Norwegian and Hungarian material, matching that of the type of C. vibratilis var. velenovskyi Rob. Henry ex Bidaud, Moënne-Locc. & Reumaux (K. Liimatainen, pers. comm.). Photo: Bálint Dima Fig. 13 — Cortinarius emollitus (Hungary), DB 1067 Habitat and distribution: In acidophilous-mesophilous Quercus(-Corylus-Tilia) forests, also found once in calcareous Tilia forest. Hitherto known with certainty (including genetic analysis) only from S Norway, S Sweden (K. Liimatainen, pers. comm.) and Hungary, but collections from SW Germany probably also belong here (Saar, Schmidt-Stohn pers. com.). Investigated material from NO and HU: NORWAY: Oslo; Bygdøy, Dronningberget, Ilkka Kytövuori, TEB 663-11 (Tilia). Vestfold; Larvik, I. Kristoffersen & S. Aasrum, TEB 689b-11 (Quercus); Larvik, TEB 300-10 (Tilia-Quercus). Aust-Agder; Arendal, TEB 652-11 (Quercus). Froland, TEB 302-09 (Quercus). Tvedestrand, TEB 637-11 (Quercus-Tilia). HUNGARY: HajdúBihar; Hosszúpályi, DB 1067 (Quercus). Pest; Budakeszi, DB 1771 (Quercus). Comments: The species resembles C. vibratilis s. l., but differs in its normally characteristic innately ibrillose cap and habitat in oak (versus coniferous) forests. The species is morphologically variable, from rather small and slender to large and robust. In Hungary almost exclusively associated with Quercus spp. and it seems to be a character species of the acidophilous oak (frondose) forests. In Norway it occurs both in poor to rich, dry oak forest types, preferentially in mesotrophic ones. However, C. emollitus is fairly overlooked and the real distribution is unknown, due to the confusion with C. vibratilis s. l. The protologue and unpublished plate of C. emollitus in Fries (FriES 1838) it well with our species; habitat indications correspond, and Fries indicates also that this has an innately ibrillose cap. 24 Acknowledgements We would like to thank the «Phlegmacium-group» organized under J.E.C. for their cooperation, including DNA analyses: Francesco Bellù, Tobias Frøslev, Bernhard Oertel, Karl Soop, Günter Saar. Furthermore, László Albert (Budapest), Egil and Katriina Bendiksen (Oslo) and Ilkka Kytövuori, Kare Liimatainen, Tuula Niskanen (Helsinki) are also thanked for providing collections, interpretations of sequences and useful information about some of the species. Norwegian data from 2010–2011 are based on fundings from the Species Project of the Norwegian Biodiversity Information Centre («Artsdatabanken»), Trondheim. Literature AlBErT, L. (1996). Cortinarius olivascentium. Olívsárga pókhálósgomba. Mikol. Közlem., Clusiana 35(3): 75. Ballarà, J. (1999). Alguns Cortinarius interessants dels Pirineus catalans. II. Rev. Cat. Micol. 22: 47–70. Bidaud, A. (2011). Cortinaires rares ou nouveaux de la région Rhône-Alpes (France). Journ. J.E.C. 13: 4–24. Bidaud, A., MoënnE-Loccoz, P. & REumaux, P. (1993). Atlas des Cortinaires. Pars V. Ed. Fédérat. Mycol. Dauphiné-Savoie, Annecy. Bidaud, A., MoënnE-Loccoz, P. & REumaux, P. (1995). Atlas des Cortinaires. Pars VII. Ed. Fédérat. Mycol. Dauphiné-Savoie, La Roche-sur-Foron. Bidaud, A., MoënnE-Loccoz, P. & REumaux, P. (1996). Atlas des Cortinaires. Pars VII. Ed. Fédérat. Mycol. Dauphiné-Savoie, La Roche-sur-Foron. BohuS, G. (1970). Interessantere Cortinarius-Arten aus dem Karpaten-Becken. Annls hist.-nat. Mus. natn. Hung. 62: 137–148. BohuS, G. (1984). Studies on the pH requirement of soil-inhabiting mushrooms: the R-spectra of mushroom assemblages in deciduous forest communities. Acta Bot. Hung. 30(1–2): 155–171. BohuS, G. & BaBoS, m. (1967). Mycocoenological investigation of acidophilous deciduous forests in Hungary. Bot. Jahrb. 87: 304–360. Brandrud, T. E. (2007). Rødlistearter av sopp knyttet til edellauvskog; habitatkrav, hotspothabitater og utbredelsesmønstre. (Redlisted species of fungi in Norway associated with thermophilous deciduous forests: habitat preferences, hotspot-habitats and distributional patterns.) Agarica 27: 91-109. Brandrud, T. E. & BEndikSEn, E. (2001). The Cortinarius species of calciphilous Tilia-Corylus and QuercusCorylus woodlands of Fennoscandia, outposts of the temperate Fagus and Quercus-Carpinus forest types of C. Europe. Journ. J.E.C. 3: 105-113 [Engl. version]. Brandrud, T. E., BEndikSEn, E., BEndikSEn, K., kyTövuori, I. & ToivonEn, M. (2012). Korallsopper (Ramaria ss. str.) i eikeskog og annen edellauvskog i Norge. (Ramaria ss. str. species in oak forests and other thermophilous, deciduous forests in Norway). Agarica 32 (in print.) Brandrud, T. E., KnuTSSon, T. & Soop, K. (1996). En fantastisk säsong på Öland. Rapport med tonvikt på Phlegmacium (spindelskivlingar). Jordstjärnan 17(3): 27–42. Brandrud, T. E., hanSSEn, O., SvErdrup-ThyGESon, A. & ødEGaard, F. (2011). Kalklindeskog – et hotspothabitat. Sluttrapport under ARKO-prosjektets periode II. (Calcareous lime forest – a hotspot-habitat. Final report from the second period of the ARKO-project) - NINA Rapport 711. Brandrud, T. E. & SchmidT-STohn, G. (2011). Der Huy – ein artenreicher Cortinarien-Standort mit Kalklaubwäldern in Sachsen-Anhalt. Journ. J.E.C. 13: 63–78. BrEiTEnBach, J. & kränzlin, F. (2000). Pilze der Schweiz. Band 5 – Agaricales III Cortinariaceae. Edition Mykologia, Lucerne. CadiñanoS AGuirrE, J. A. (2006). Algunos Sericeocybe del grupo del Cortinarius alboviolaceus (subsección Sericeocybe Bidaud et al.) recolectados en el norte de España. Journ. J.E.C. 8: 3–42. ConSiGlio, G. (1997). Contributo alla conoscenza dei macromiceti della regione Emilia-Romagna. VII. Genere Cortinarius – Parte quarta. Riv. Micol. 40: 159–174. 25 EySSarTiEr, G. (2004). Notes sur quelques espèces de cortinaires et de psathyrelles rares ou nouvelles. Fungi Non Delineati 28. Edizioni Candusso, Alassio, 55 pp. dähnckE, R. M. & dähnckE, S. M. (1979). 700 Pilze in Farbfotos. AT Verlag, Stuttgart. FriES, E. M. (1838). Epicrisis Systematis mycologici. Upsaliae. FrøSlEv, T. G., JEppESEn, T. S., læSSøE, T. & kJøllEr, R. (2007). Molecular phylogenetics and delimitation of species in Cortinarius section Calochroi (Basidiomycota, Agaricales) in Europe. Molecular Phylogenetics and Evolution 44: 217–227. FrøSlEv, T. G., maThEny, p. B. & hiBBETT, d. S. (2005). Lower level relationships in the mushroom genus Cortinarius (Basidiomycota, Agaricales): A comparison of RBP1, RBP2, and ITS phylogenies. Molecular Phylogenetics and Evolution 37: 602–618. hEnry, r. (1951). Les Scauri. Bull. Soc. mycol. France 67: 225–322. HEnry, R. (1966). Étude ou révision de quelques Cortinaires appartenant aux Myxacia et aux Scauri. Bull. Soc. mycol. France 82: 135–178. HEnry, R. (1981). Les Cortinaires. Bull. Soc. mycol. Fr. 97(3): 159–334. JEppESEn, T. S. & FrøSlEv, T. G. (2011). Knoldslørhatte som indikatorarter gennem 20 år (1991-2011). Svampe 64: 34-45. knudSEn, H. & vESTErholT, J. (2008) Funga Nordica. Agaricoid, boletoid and cyphelloid genera. Nordsvamp. Copenhagen. kålåS, J. a., vikEn, å., hEnrikSEn, S. & SkJElSETh, S. (eds.) (2010). The 2010 Norwegian Red List for Species. Norwegian Biodiversity Information Centre. Trondheim. KriEGlSTEinEr, G. J. (1991). Verbreitungsatlas der Großpilze Deutschlands (West). Band 1: Ständerpilze. Teil B: Blätterpilze. Ulmer, Stuttgart. GmindEr, A. & KriEGlSTEinEr, G. J. (2010). Die Großpilze Baden-Württembergs. Band 5: Ständerpilze: Blätterpilze III. Ulmer, Stuttgart. PödEr, R. (1982). Contributo ed ossevazioni sul Symposium internazionale e mostra di micologia. Atti Symp. intern. micol. Borgotaro 1: 155–182. rimóczi, I. & alBErT, L. (1992): Cortinarius-Arten in Ungarn. Beitr. Kenntn. d. Pilze Mitteleur. 8: 208–220. Saar, G. (2010). Die Arten aus dem Cortinarius largus-/variecolor-Komplex, Sektion Phlegmacioides (= Variecolores). Journ. J.E.C. 12: 42–56. SchmidT-STohn, G. & oErTEl, B. (2009). DNA-Analysen in der Pilz-Taxonomie. Journ. J.E.C. 11: 10-19. vaSaS, G. & locSmándi, cS. (2009). The Basidiomycetes of the Aggtelek National Park. In: papp B. (ed.): Flora of the Aggtelek National Park, Cryptogams. Hungarian Natural History Museum, Budapest, pp. 53–107. VEllinGa, E. C. (1986). Contribution à la connaissance de la lore mycologique de la Haute Belgique, cinquème inventaire. Nat. Mosana 39(3): 63–80. 26