Cortinarius subgenus Phlegmacium section Multiformes in Europe

Balint Dima; Karl Soop; Tobias Frøslev; Geert Schmidt-stohn; Günter Saar

The taxonomy and phylogeny of sect. Multiformes (Cortinarius, subgen. Phlegmacium) are presented. The present circumscription of the section is supported by molecular data and morphological features. The taxonomy is complex and many taxa are quite new and little known. Two new species are described. Altogether ten European species are recognized: C. multiformis, C. talimultiformis, C. frondosomultiformis sp. nov., C. rufoallutus, C. talus, C. melleicarneus, C. caesiolamellatus, C. caesiophylloides, C. pallidirimosus and C. armenicorius sp. nov. The ten recognized species are (i) more or less well-supported phylogenetically, but (ii) with a fairly low morphological differentiation. The species differs by >1% in ITS DNA from sister taxa, and each species pair show a morphological differentiation, though in most cases with overlapping characters. For instance, most species have overlapping variation in size and shape of the spores, and less than half of the species have formerly been recognized in morphological studies.

Journal des J.E.C. No. 16., p. 162 – 199, 2014 Cortinarius subgenus Phlegmacium section Multiformes in Europe Tor Erik Brandrud1*, BálinT dima**, GEErT SchmidT-STohn*** FrancESco BEllù****, ToBiaS G. FrøSlEv*****, BErnhard oErTEl****** GünTEr Saar******* & karl Soop********* received on 19 August 2014 Summary The taxonomy and phylogeny of sect. Multiformes (Cortinarius, subgen. Phlegmacium) are presented. The present circumscription of the section is supported by molecular data and morphological features. The taxonomy is complex and many taxa are quite new and little known. Two new species are described. Altogether ten European species are recognized: C. multiformis, C. talimultiformis, C. frondosomultiformis sp. nov., C. rufoallutus, C. talus, C. melleicarneus, C. caesiolamellatus, C. caesiophylloides, C. pallidirimosus and C. armenicorius sp. nov. The ten recognized species are (i) more or less well-supported phylogenetically, but (ii) with a fairly low morphological differentiation. The species differs by >1% in ITS DNA from sister taxa, and each species pair show a morphological differentiation, though in most cases with overlapping characters. For instance, most species have overlapping variation in size and shape of the spores, and less than half of the species have formerly been recognized in morphological studies. ZuSammenfaSSung Es wird die Taxonomie und Phylogenie der Sekt. Multiformes (Cortinarius, Untergattung Phlegmacium) vorgestellt. Die gegenwärtige Abgrenzung der Sektion wird durch molekulare Daten und morphologische Eigenschaften gestützt. Die Taxonomie ist kompliziert, und viele Arten sind ganz neu und wenig bekannt. Zusammen 10 europäische Arten sind anerkannt: C. multiformis, C. talimultiformis, C. frondosomultiformis sp. nov., C. rufoallutus, C. talus, C. melleocarneus, C. caesiolamellatus, C. caesiophylloides, C. pallidirimosus und C. armenicorius sp. nov. Diese 10 Arten sind (i) phylogenetisch mehr oder wenig gut gesichert, allerdings (ii) morphologisch ziemlich wenig differenziert. Die Arten unterscheiden sich in >1% ihrer ITS-DNA von den Schwester-Arten, und jedes Arten-Paar zeigt zwar eine morphologische Differenzierung, die sich aber meist überlappt. So sind die Sporenmaße und die Sporenform der meisten Arten recht * Tor Erik Brandrud, Norwegian institute for nature research (NINA), Gaustadalléen 21, 0349 Oslo, Norway; tor.brandrud@nina.no ** Bálint Dima, Department of Biosciences, Plant Biology, University of Helsinki, P.O. Box 65, 00014 Helsinki, Finland; cortinarius1@gmail.com *** Geert Schmidt-Stohn, Burgstraße 25, D-29553 Bienenbüttel, Germany; geert.schmidt-stohn@t-online.de **** Francesco Bellù, CP 104, I-39100 Bolzano, Italy; bellu.francesco@rolmail.net ***** Tobias G. Frøslev, Natural History Museum of Denmark, Center for Geogenetics, University of Copenhagen, Øster Voldgade 5–7, 1350 København K, Denmark; tobias.froeslev@gmail.com ****** Bernhard Oertel, INRES, Universität Bonn, Auf dem Hügel 6, D-53121 Bonn; b.oertel@uni-bonn.de ******* Günter Saar, Dammenmühle 7, D-77933 Lahr-Sulz, saar.guenter@t-online.de ******** Karl Soop, Honorary Research Associate, Swedish Museum of Natural History, Dept. of Cryptogamic Botany, Djustäppv. 23, S-792 90 Sollerön, Sweden; karl@soop.org 162 ähnlich, und weniger als die Hälfte der hier vorgestellten Arten wurden früher in morphologischen Studien unterschieden. réSumé On présente la taxinomie et la phylogénie de la section Multiformes (Cortinarius, sous-genre Phlegmacium). La délimitation actuelle de la section est étayée par des données moléculaires et des caractères morphologiques. La taxinomie est compliquée et beaucoup d’espèces sont tout à fait nouvelles et peu connues. On reconnait en tout 10 espèces européennes : C. multiformis, C. talimultiformis, C. frondosomultiformis sp. nov., C. rufoallutus, C. talus, C. melleocarneus, C. caesiolamellatus, C. caesiophylloides, C. pallidirimosus et C. armenicorius sp. nov. Ces 10 espèces sont (i) plus ou moins bien identiiées au plan phylogénétique, toutefois (ii) assez peu différenciées morphologiquement. Les espèces diffèrent des espèces soeurs à raison de >1% quant à leur ITS d’ADN, et chaque paire d’espèces présente certes une différenciation morphologique, mais avec des chevauchements. C’est ainsi que les dimensions et la forme des spores de la plupart des espèces sont très semblables, et moins de la moitié des espèces présentées ici étaient autrefois distinguées dans les études morphologiques. riaSSunto Viene presentata la tassonomia e la ilogenetica della sezione Multiformes (Cortinarius, sottogenere Phlegmacium). L’attuale distinzione della sezione si basa su dati molecolari e caratteri morfologici. La tassonomia è complicata e molte specie sono completamente nuove e poco note. Complessivamente sono riconosciute 10 specie europee: C. multiformis, C. talimultiformis, C. frondosomultiformis sp. nov., C. rufoallutus, C. talus, C. melleocarneus, C. caesiolamellatus, C. caesiophylloides, C. pallidirimosus and C. armenicorius sp. nov. Queste 10 specie sono supportate più o meno bene da un punto di vista ilogenetico (i), ma (ii) differenziate abbastanza poco da un punto di vista morfologico. In >1% del loro ITS-DNA, le specie si distinguono dai taxa sorella ed ogni copia di specie permette una differenzazione morfologica, però con frequenti sovrapposizioni. Pertanto, dimensioni e forma delle spore sono molto simili tra di loro, e meno della metà delle specie qui presentate venivano in passato distinte attraverso studi morfologici. reSumen Presentamos la taxonomía y la ilogenia de la secc. Multiformes (Cortinarius, subgén. Phlegmacium). La presente circunscripción de la sección está basada en los resultados moleculares y en las características morfológicas. La taxonomía es compleja por lo que muchos taxones son totalmente nuevos y poco conocidos. Se describen dos nuevas especies. En total se reconocen diez especies europeas: C. multiformis, C. talimultiformis, C. frondosomultiformis sp. nov., C. rufoallutus, C. talus, C. melleicarneus, C. caesiolamellatus, C. caesiophylloides, C. pallidirimosus and C. armenicorius sp. nov. Las diez especies reconocidas lo son (i) con más o menos buen soporte ilogenético, pero (ii) con una diferenciación morfológica bastante baja. Las especies diieren por >1% en ITS DNA de los taxones más cercanos, y cada par de especies muestra una diferenciación morfológica, aunque en muchos casos haya una superposición de caracteres. Por ejemplo: muchas especies muestran una variación en la superposición del tamaño y la forma de las esporas, y sin embargo menos de la mitad de las especies han sido diferenciadas anteriormente mediante estudios morfológicos. 163 Table 1. Collections used for phylogenetic analysis. Species are presented in alphabetical order. Species C. armenicorius sp. nov. C. armenicorius C. caesiolamellatus C. caesiolamellatus Collector* KS UP UL SSt, TEB Locality France, Jura, Prénovel Herbarium KS-CO1865 (Holotypus) IB19990590 TUB011841 TEB428-10 GenBank KM504515 AF325586 AY669531 unpublished TEB687-11 unpublished C. rufoallutus var. caesiolamellatus C. multiformis var. caesiophyllus C. caesiophylloides AB Russia, Sakha, Khangalassky Ulus Germany, Thürringen, Thölendorf Baden-Württemberg, Schwaben, Ehingen a.d. Donau, Kohlhau Norway, Oslo, Grorud, Steinbruvannet Norway, Oslo, Vestfold, Tønsberg Norway, Sogn & Fjordane, Lærdal Italy, Malga di Nemes USA, Washington, Olympic peninsula France, Ain, col des Bérentin PML France, Savoie, Arith PML882 (Holotypus) KF732351 TEB TEB277-09 unpublished C. caesiophylloides C. caesiophylloides TF TEB Norway, Nord-Trøndelag, Stjørdal, Beistadvollen Norway, Oppland, Lunner Norway, Buskerud, Ringerike, Eggemoen northeast Finland, ES, Joutsa, Koivuranta Finland, Kn, Paltamo Italy, South Tyrol, Kaltern east, Montigglerwald near Eppan Italy, South Tyrol, Kaltern west, Matschatsch (route to Mendelpass) Italy, South Tyrol, Kaltern west, Matschatsch (route to Mendelpass) Estonia, Hiiumaa, Sarve, Soonlepa Norway, Aust Agder, Grimstad Sweden, Dalarna, Revnes Italy, South Tyrol, Sarntal, Reinswald Norway, Oppland, Øystre Slidre, Reieåsen Norway, Oppland, Vestre Slidre, Mosåni Norway, Oppland, Øystre Slidre, Volbufjorden Norway, Hordaland, Voss Austria, Zillertal Sweden, Jämtland, Ragunda, Bispgården Austria, Gepatsch-Stausee Italy, Corti, Grünbachalm Italy, Pochi di Salorno, Val Fredda Sweden, Borgsjö/Jämtgaveln Estonia, Võru, Rõuge, Saarlase Sweden, Ångermanland, Häggdånger, Sjö TF2006-112 DB-NO-181 TEB720-13 TN05-016 (Holotypus) IK08-1554 TG2000-218 (Holotypus) TF2000-086 KF732575 unpublished Bellu07-11-2011 (BOZ 2011/71) IK01-053 (Holotypus) O125960 KS-CO1974 Bellu21-09-2011 unpublished DB-NO-33 TEB410-13 TEB186-07 unpublished DB-NO-37 TEB413-13 TN05-247 MD12-2012 AT2004187 unpublished KF732626 unpublished UDB002163 SSt09-017 TG1999-231 TG1998-096 SSt10-068 TU105180 CFP445 (Neotypus) unpublished unpublished unpublished unpublished UDB016130 KF732350 C. caesiolamellatus TEB C. caesiolamellatus C. caesiolamellatus C. caesiolamellatus C. caesiolamellatus TEB TEB CR JFA, TN C. caesiophylloides C. caesiophylloides C. frondosomultiformis sp. nov. C. frondosomultiformis KL, TN IK GT C. frondosomultiformis AC C. melleicarneus C. melleicarneus C. multiformis C. multiformis IK ILF KS FB TF, TSJ C. multiformis TEB, BD C. multiformis TEB C. multiformis TEB, BD C. multiformis C. multiformis C. multiformis TN et al. MD AT C. multiformis C. multiformis C. multiformis C. multiformis C. multiformis C. multiformis SSt GT GT SSt AK TEB et al. 164 TEB278-80 TEB213-12 Rossi05-08-012 TN09-201 unpublished unpublished unpublished KF732571 PML4905 (Holotypus) KF732414 KF732572 KF732574 KM504516 unpublished KF732577 AY669533 unpublished unpublished unpublished Species C. multiformis C. pallidirimosus C. pallidirimosus C. pallidirimosus C. pallidirimosus C. pallidirimosus C. rufoallutus C. rufoallutus C. rufoallutus C. rufoallutus C. rufoallutus C. sp. C. sp. C. talimultiformis C. talimultiformis C. talimultiformis C. talimultiformis C. aurantionapus var. similis C. talimultiformis C. talimultiformis C. talimultiformis C. talimultiformis C. talimultiformis C. talimultiformis C. talus C. talus C. talus C. pseudotalus C. ochropudorinus C. pseudominor C. crenulatus C. talus C. talus C. talus C. talus C. talus C. calochrous (outgroup) Collector* TN et al. IK IK TN, KL IK IK CR TEB, BD TEB PML, JM AK CR SSt GT SSt PML AT IK LA, BD KP GS TEB et al. AK JCM AB PR FL TEB, SSt SSt, TEB TF KT TF Locality Herbarium Finland, PK, Kitee TN06-139 Finland, InL, Utsjoki, Kevo IK95-585 (Holotypus) Finland, PeP, Tervola IK07-692 (H6000925) Finland, PeP, Rovaniemi TN04-470 Finland, SoL, Sodankylä IK92-966 Norway, Troms, Storfjord IK98-711 Italy, South Tyrol, Anterselva Rossi02-08-2006 Norway, Oppland, Øystre Slidre, DB-NO-38 Volbufjorden TEB 415-13 Norway, Oppland, Øystre Slidre TEB460-11 France, Haute-Savoie, Plateau de PML635 (Holotypus) Glières Sweden, Jämtland, Borgsjö AV01-09-97 Estonia, Hiiu, Pühalepa, Salinõmme TAAM128770 Germany TUB019739 Italy, Tovare Male’ Rossi29-08-2002 Germany, Baden-Württemberg, SSt10-188 Ehingen a.d. Donau Italy, Brunico/Amaten TG1999-246 Germany, Niedersachsen, Ahlsburg/ SSt09-040 Dörrigsen France, Haute-Savoie, Avernioz PML883 (Holotypus) Sweden, Uppsala, Hässelby Park Finland, PH, Virrat Turkey, Trabzon, Macka Hungary, Vas, Felsőszölnök Estonia, Tartu, Nõo, Vellavere Germany, Baden-Württemberg, Titisee-Neustadt USA Sweden, Jämtland, Ragunda Estonia, Tartu, Vara, Alajõe France France, Ain, Meyriat France, Ardennes, Bois des Alleux France, Allier, Forêt de Tronçais Sweden, Borgsjö Sweden, Borgsjö Germany Denmark, Sealand Hungary, Vas, Farkasfa Sweden, Skåne GenBank KF732624 KF732578 KF732579 KF732580 KF732581 KF732582 unpublished unpublished unpublished KF732413 KF732639 UDB015964 unpublished unpublished unpublished unpublished unpublished KF732257 AT2004096 (Holotypus) H6032747 SES2741 UDB001167 KF732584 KF732585 DB2831 TAAM128693 TUB0118410 unpublished UDB015959 AY669532 SAT00-261 CFP832 (Neotypus) TU105173 PML4859 (Holotypus) PML2339 (Holotypus) PML4750 (Holotypus) PML4866 (Holotypus) TEB140-10 SSt10-033 TUB019737 TF1999-035 DB4435 TF2001-113 FJ717579 KF732457 UDB016004 KF732402 KF732361 KF732396 KF732292 unpublished unpublished unpublished unpublished unpublished DQ663250 * Abbreviation of names: AB = André Bidaud, AC = A. Cester, AK = Anu Kollom, AT = Andy Taylor, BD, DB = Bálint Dima, CR = Claudio Rossi, FB = Francesco Bellù, FL = F. Lopez, GS = Günter Saar, GSS = Geert Schmidt-Stohn, GT = Gianni Turrini, ILF = I-L. Fonneland, IK = Ilkka Kytövuori, JCM = J.-C. Malaval, JFA = Joseph F. Ammirati, JM = Jacques Melot, KL = Kare Liimatainen, KP = Kai Piirsoo, KT = Katalin Takács, LA = László Albert, MD = Martin Dondl, PML = Pierre Moënne-Loccoz, PR = Patrick Reumaux, TEB = Tor Erik Brandrud, TF = Tobias G. Frøslev, TM = Thomas Münzmay, TN = Tuula Niskanen, TSJ = Thomas Stjernegaard Jeppesen, UL = Udo Luhmann, UP = Ursula Peintner. 165 Phylogram: Bálint Dima Fig. 1 – Maximum likelihood ITS phylogram of Cortinarius sect. Multiformes in Europe calculated with RAxML. Bootstrap values are given above branches. The bar indicates the number of expected substitutions per position. For each collection, herbarium number, locality (country), and GenBank/UNITE accession numbers if existed are provided. 166 Illustration: Marie K. Brandrud Fig. 2 – Schematic illustration of the morphological, ecological and phylogenetic afinity among the sect. Multiformes species. Circles: deciduous forest species. Squares: coniferous forest species. thick line: preferentially northern (middle- northern boreal) species. Thin line: more southern species (mainly S. Scandinavia-C Europe). Green: species with pale yellowish (whitish to honey brown) pileus. Red: species with vivid ochraceous brown to red brown pileus. Blue: species with violaceous blue tinges on young lamellae and stipe. Double line between taxa = sister species with a high morphological similarity (overlap in character variation). 167 Introduction The section Multiformes (Rob. Henry) Moënne-Locc. & Reumaux (Cortinarius, subgen. Phlegmacium) is here treated in a strict sense, including species with (i) a more or less abruptly bulbous stipe, (ii) context with a honey-like smell, (iii) a duplex pileipellis structure and (iv) subamygdaloid-ellipsoid, pale, rather densely verrucose spores. This group, in contrast to the resembling sect. Claricolores treated in Brandrud et al. (2013), consists of a number of little known, critical species, and its taxonomy has not been well-established. According to morphological taxonomy, only two or three species have frequently been distinguished up to now; the ochraceous (orange-)brown C. multiformis (sometimes also C. allutus or C. rufoallutus) in coniferous forests and the paler C. talus (= C. ochropallidus, C. multiformis s. Moser) in deciduous forests (cfr. e.g. Brandrud et al. 1990–2014, JEppESEn et al. 2012, kriEGlSTEinEr & GmindEr 2010, Soop 2011). MoSEr (1961) distinguished three species in coniferous forests referable to the present C. multiformis (C. allutus, C. lundellii and C. subhygrophanicus) and three in deciduous forests referable to C. talus as applied here (C. multiformis, C. ochropallidus and C. talus). However, recent and ongoing molecular studies performed by the present authors in the JEC DNA group, by the Tübingen group (Garnica et al. 2014 in prep.) and by the Helsinki group (liimaTainEn et al. 2014) have shown that the taxonomy of sect. Multiformes is much more complex than formerly thought. With this new, phylogenetic knowledge, it has become impossible to maintain the old, wide species concept based on discontinuous morphological variation. The sect. Multiformes has often been treated in a wider concept, including most phlegmacioid species with whitish-greyish lamellae and context. For instance, species with a more cylindricclavate stipe, a duplex pileipellis and amygdaloid-ellipsoid spores, covering the C. claricolor-C. cumatilis-C. praestans group (sect. Claricolores) as well as the C. turmalis-C. variegatus group has been included here (see e.g. Brandrud et al. 1990–2014). However, Garnica et al. (2005, 2014 in prep.) have shown that sect. Claricolores («clade Praestantes») and the C. turmalis-C. variegatus group are phylogenetically rather dissimilar from sect. Multiformes, especially the former, which appears to be more closely related to sect. Phlegmacioides and to sect./clade Vulpini (cfr. also Brandrud et al. 2013). Phylogenetically, sect. Multiformes s. str. as applied here appears to be a well supported clade (see Garnica et al. 2005; sub nom. clade Alluti). Species such as C. saporatus and C. corrosus with (i) whitish-greyish lamellae and context, (ii) a strongly marginate bulbous stipe, (iii) a simplex, thick pileipellis and (iv) and strongly-coarsely ormamented citriform-amygdaloid spores have also often been included in (sub)sect. Multiformes (see e.g. MoSEr 1961, Bidaud et al. 2006), but these are clearly both micromorphologically and phylogenetically quite dissimilar from the «true» Multiformes species, and belong to the well-circumscribed sect. Calochroi (cfr. e.g. FrøSlEv et al. 2007, Garnica et al. 2005). Material and Methods The presented species have been studied from various parts of Europe, including documentations with photographs and descriptions of macro- and micromorphology. The study of macroand micromorphology follows Brandrud (1996). For spore measurements, unripe or aberrant spores are avoided, and only spore deposits (including spores on the stipe/Cortina) are measured. Spore size variation as shown in spore-scattergrams is based on mean values per basidiome. One or two basidiomes are measured per collection. For method of spore photos with stacking techniques, see SchmidT-STohn (2011). A selected set of samples were also used for phylogenetic studies, combined with already available sequences from the online DNA-sequence depositories GenBank (http://www.ncbi.nlm.nih.gov/gen-bank/) and UNITE (http://unite.ut.ee/). Laboratory work was organized/performed by some member of the JEC DNA group (Geert Schmidt-Stohn, Tobias G. Frøslev, Tor Erik Brandrud, Bálint Dima) in the following universi- 168 ties: Greifswald and Senckenberg (Germany), Copenhagen (Denmark), Oslo (Norway) and in the Eötvös Loránd University, Budapest (Hungary). Main responsible for the laboratory work at the University of Greifswald was Martin Unterseher, and at the research centre Senckenberg (Frankfurt a. M.) Ali Tahir and Marco Thines. For the DNA extraction and PCR reactions mainly the Phire® Plant Direct PCR Kit (Finnzymes, Finland) was used, but also other DNA extraction kits were applied (see FrøSlEv et al. 2005, 2007, cfr. also SchmidT-STohn & OErTEl 2009). The ITS region of the nrDNA was ampliied with the primer pairs ITS1/ITS4, ITS1F/ITS4 or ITS1F/ITS4B. Majority of PCR products were sequenced by LGC Genomics (Berlin, Germany) with the same primers used in PCR reactions. The electropherograms were checked, assembled and edited with the CodonCodeAligner 4.1. (CodonCode Corporation, Centerville, MA, U.S.A). Sequences produced by the JEC-DNA Group are unpublished; the data are available from the second author. Alignment was performed by webPRANK (LöyTynoja & Goldman 2010, http:// www.ebi.ac.uk/goldman-srv/webPRANK/). Maximum Likelihood analysis was carried out with RAxML BlackBox (STamaTakiS et al. 2008, http://embnet.vital-it.ch/raxml-bb/). Cortinarius calochrous (DQ663250, FrøSlEv et al. 2007) was chosen as outgroup. Altogether 72 sequences (+ outgroup) were included in our RAxML analysis of which 34 were generated in this study (Fig. 1). Many of our sequenced collections were, however, excluded from the analysis due to their insuficient length or quality. Nevertheless, these sequences could be used for identiication using BLAST alogorithm of GenBank and/or UNITE databases. All collections examined morphologically by us are listed under each species description. The sequenced collections (own and public databases) are marked (DNA), and those included in the phylogenetic analysis are marked with an additional asterisk (DNA*). Section Multiformes – 10 species recognized in Europe As presented and discussed below, the phylogenetic data provides support for a considerably number of new, formerly little known taxa in sect. Multiformes. Four of these species are described as new in LiimaTainEn et al. (2014), and two in the present study. In some cases, close taxa are more or less geographically differentiated. For instance in the C. multiformis complex, the more Northern European, oligotrophic-acidophilous C. multiformis s. str. (= C. lundelli (M. M. Moser) M. M. Moser) can be distinguished from the apparently more Central-Southern European C. talimultiformis (= C. aurantionapus var. similis Moënne-Locc.), while showing overlapping morphological features. The blue gilled species show a similar differentiation; C. caesiophylloides appears to be a predominantly Northern European, middle boreal-northern boreal taxon, whereas the closely related C. caesiolamellatus seems to be predominantly South ScandinavianCentral(-South) European, southern boreal-montane. On re-examination of the morphology of phylogenetic units, it appears that some degree of morphological differentiation can be found between all taxa, but some taxa will be dificult to unambiguously identify in the ield or under the microscope. In fact, it is not possible to fully understand the taxonomy of sect. Multiformes without the aid of molecular characters and phylogeny. This is partly due to (i) the low degree of morphological differentiation, with sister species showing overlap in morphological characters, but also due to (ii) the rarity of some of the taxa. For instance, C. melleicarneus is hitherto found twice, once in Estonia (2001) and once in southernmost Norway (1994), and C. armenicorius is also found only twice; once in Russia (1999) and once in French Jura (2008). Cortinarius frondosomultiformis is only found in two sites in N Italy (2000, 2011). It will probably take more than a lifetime to get good data on the morphological variation and ecogeographical range of such rarities, and sequence data are thus essential. This taxonomy of Multiformes is comparable to that of some critical groups of subgenus Telamonia, where many taxa are well-supported phylogenetically, but with comparatively little 169 morphological differentiation (e.g. sect. Brunnei, NiSkanEn et al. 2009, sect. Bovini niSkanEn et al. 2013). These seem to represent young groups in active speciation, and introducing molecular methods has resulted in (more than) a doubling of the number of species. In other groups of subgenus Phlegmacium, such as sect. Claricolores (Brandrud et al. 2013), sect. Phlegmacium (cfr. Brandrud 1996 versus Garnica et al. 2005) and some fulvoid groups (e.g. the Rufoolivacei) of sect. Calochroi (Brandrud et al. in prep.) the new, molecular data has hardly changed the wide, well-established morphology-based species concepts formerly applied. Since the new combined phylogenetic and morphological species concept support the inclusion of a number of morphologically relatively weakly differentiated species that sometimes might even be dificult to distinguish morphologically, this might be taken as an argument for a general, narrow morphological species concept. However, the phylogenetic results do not support the former «splitting taxonomy» which has been perfomed mainly in France. For instance, Bidaud et al. (2006) included at least 16 species in sect. Multiformes s. str., but these apparently do not cover more than 4 of the species accepted here. So 75% of the species names applied at present in sect. Multiformes s. str. in France are synonyms or probable synonyms. For instance, at least 9 of the species distinguished in Bidaud et al. (2006), appears to be within the rather wide morphological variation of C. talus. Morphology and delimitation towards related groups The species of sect. Multiformes are characterized macromorphologically by a more or less abruptly bulbous stipe with only little universal veil remnants at the bulb margin. Veil remnants are sparse also on the pileus, but some thin, silvery white veil hyphae might give the surface a frosty or marbled-innately ibrillose pattern on young basidiomes. Cortina remnants are also thin, often rendering the stipe surface smooth and glossy, initially being whitish(-bluish) and often becoming shiny brass brown with age. The pileus is often bicoloured, that is, darker hygrophanous (water-soaked) in outer part, at least with age and when growing in humid habitats, or the surface may be pseudohygrophanous, that is with a dense pattern of darker spots at centre and hygrophanous streaks or veins towards margin. Some species become radially rugulose-sulcate (wrinkled) with age due to the cap cuticle structure. Micromorphologically, the following features are characteristic; (i) a duplex (two-layered) pileipellis structure including a subcellular hypoderm with tightly cemented elements provided with golden yellow, parietal, amber-like pigment (usually without spirally-zebra-striped encrustations), and (ii) the rather pale, amygdaloid-ellipsoid spores with somewhat rounded warts. The spore ornamentation usually consists of irregular, low, rounded and often conluent warts, which might appear rather diffuse in the microscope (Figure 3, 12). The spore morphology is rather uniform in the section, however with some small differences in size and shape that can be used to more or less distinguish closely related taxa. One «series» of species with rather strictly(acutely) amygdaloid spores can be distinguished from species with more broadly ellipsoid spores (Figure 3, 12). The former group has slightly narrower spores, with average Q-values ranging from 1.68–1,81, whereas the ones with more broadly ellipsoid spores possess mean Q values of 1.60–1.71. Normally, the strictly amygdaloid spores are paler, with dense ornamentation, whereas the more ellipsoid spores are darker (more thick-walled) with more coarse, conluent ornamentation (Figure 1). For instance, the close species pair C. multiformis versus C. talimultiformis can be distinguished on more broadly ellipsoid, coarsely verrucose spores versus strictly amygdaloid more densely verrucose and pale spores, respectively. The same applies for C. caesiolamellatus versus C. caesiophylloides. However, it should be noted that the spore morphology variation might overlap, for instance, species with mainly ellipsoid spores might sometimes have dominating amygdaloid spores. Mature spores shed to the stipe or in spore deposits should always be compared for these patterns. 170 According to material seen so far, C. frondosomultiformis have smaller spores than the others (7‒8.5 × 4.5‒5 µm versus «standard» Multiformes spores 8‒9.5 × 5‒5.5 µm), and also C. melleicarneus has slightly smaller spores than the average Multiformes species (8‒8.5 × 4.5‒5 µm) (Figure 3, 12). On the other hand, C. armenicorius, C. pallidirimosus and C. talimultiformis possess slightly larger spores than average. Macrochemical reactions vary considerably within the species (dependent on age, weather conditions, etc.) and are regarded as of limited taxonomic value, so is not included here. Chemically, the section Multiformes taxa are characterized by a strong, bluish luorescent substance (possibly a sterol). This can be observed on dried specimens under UV light, and is pronounced when tissue is extracted and run by chromatographical methods (Brandrud 1980). The sect. Multiformes species share a number of the above mentioned features with the members of some other, morphologically resembling taxa, such as sect. Claricolores and the C. turmalis-C. variegatus group. Section Claricolores, however, does not belong to the same major clade as sect. Multiformes, and the species of the former are distinguished by their thick, almost membranous veil, rendering the stem more or less volvate or repeatedly girdled, and the cap often with whitish patches when young. Furthermore, the species never possess an abruptly bulbous stipe, they never smell of honey, and their spores are acutely amygdaloid with little/no ornamentation or much larger (Brandrud et al. 2013). The C. turmalis-C. variegatus group belongs to the same major clade as the Multiformes group, but is not in a sister relationship, and differs by small, smooth, very pale brown spores, persistently rather pale lamellae and veil/basal mycelium which turns pink on exposure to air (cfr. Brandrud et al. 1990–2014). Molecular phylogeny A number of taxa appear to be well supported phylogenetically as shown by a study of the barcoding marker ITS, with (i) a clear barcoding gap (discontinuous sequence variation) between sister taxa, including (ii) an average ITS sequence difference between sister species of >1% (<99% similarity: (6–)8–15 nucleotide differences; see Figure 1). The C. pallidirimosus complex differs from this pattern; possessing a rather large internal ITS variation of six nucleotide differences (Figure 1). At least most of this variation is apparently not relected in corresponding morphological-ecological variation, and seems to represent infraspeciic variation. These variants, however, needs further study. As stated above, a number of sister species are morphologically very similar, especially the taxa around C. multiformis. One of the genotypes/OTU’s around C. multiformis («C. sp.») with 5–6 ITS differences from C. multiformis is not treated here due to lack of morphological data. The phylogenetic, morphological and ecogeographical afinities of the ten European Multiformes species recognized here are summarized in Figure 2. The circumpolar species C. multiformis and C. talus show 1–2 ITS sequence difference between European populations and North American populations (data not shown). The many European specimens of C. talus studied show genetically uniformity, even though the taxon is rather variable as to spore-morphology and eco-geography (see under C. talus). The extensive ITS-barcoding analyses performed lately by us and others (see e.g. Garnica et al. 2003, 2005, 2014 in prep., LiimaTainEn et al. 2014) give a clear support for the delimitation of the clade/section Multiformes (= clade Alluti) as applied here, but show conlicting and not wellsupported results for infrageneric afinity and phylogeny. However, yet unpublished data using other gene markers (Garnica et al. 2014 in prep., Høiland et al. 2014 in prep.) indicate that sect./ clade Multiformes is most closely related to the sect./clade Scauri and Purpurascentes, as well as to some smaller clades containing the European taxa C. lustratus, C. pinophilus, C. turmalis, and C. variegatus. See separate paper on sect. Purpurascentes in present volume of Journ. J.E.C. The phylogeny of these phlegmacioid groups should be regarded as not yet fully resolved. 171 Taxonomy Key to the European species of sect. Multiformes Note: The differentiation in macrocharacters is little or little known between a number of closely related species. Therefore, microcharacters (spore characters) are emphasized in the key. The often occurring honeysmell is most pronounced in bulb context. 1 1* 2 2* 3 3* 4 4* In deciduous forests. Pileus when young pale ochraceous yellow (-honey brown) to almost whitish (if more ochre brown, then spores small, MV < 8.5 µm) ....................................... 2 In coniferous (mainly Picea) forests. Pileus when young darker ochraceous brown-fulvous to vividly apricot-red brown ................................................................................................ 5 Pileus initially silvery-frosty at centre, distinctly innately ibrillose towards margin; almost whitish, later (pale) ochraceous yellow; sometimes more ochre brown. Robust species with a distinct marginate bulb. Smell strong of honey. Spores variable, (7‒)7.5–9.5 × 4.5–5.5 µm; northern, subalpine Betula pubescens populations have small spores (normally <8.5 µm long). Widespread, mainly under Betula, Fagus, Quercus, Carpinus, Corylus ....... C. talus Pileus not so frosty-innately ibrillose (if innately ibrillose, then northern, with slender stipe, and with larger spores) ............................................................................................... 3 Spores relatively large, 9–10(‒10.5) × 5–6 µm, amygdaloid; in northern Fennoscandian (boreal/subalpine) mixed Betula pubescens forests. Pileus initially (very) pale and somewhat innately ibrillose. Basidiomes slender, bulb indistinct; smell of honey .. C. pallidirimosus Spores small, 7–8.5 × 4–5 µm; in thermophilous Quercus-Corylus-Fagus forests; pileus weakly innately ibrillose or not; basidiomes robust, with a distinctly marginate bulb ...... 4 Spores narrowly amygdaloid, (7.5‒)8–8.5(‒9) × (4‒)4.5–5 µm; pileus pale honey brown to more grey brown or lesh-coloured; no smell of honey noted. In Quercus-Corylus forests on calcareous, sandy soil. Very rare (S Norway, Estonia) .............................. C. melleicarneus Spores ellipsoid-subamygdaloid, 7‒8.5(‒9) × 4‒5(‒5.5) µm, with often somewhat coarser, conluent warts; pileus ochraceous yellow to pale ochraceous brown, smell of honey. In rich Quercus-Carpinus-Fagus(-Abies) forests. Very rare (N Italy) ....... C. frondosomultiformis 5 5* Lamellae and stipe (apex) with bluish tinges when young; strong smell of honey ............. 6 Without any bluish tinges; honey smell almost absent to (rarely) strong ........................... 7 6 Spores usually ellipsoid and rather strongly and coarsely verrucose, 8–9(–9.5) × 5–6 µm. Pileus vividly, warm ochraceous brown to red brown, often bicoloured with a dark, hygrophanous outer zone. Mainly in S Scandinavia and C Europe ................ C. caesiolamellatus Spores distinctly (acutely) amygdaloid, weakly and densely verrucose, 8.5–9.5(‒10) × (4.5–) 5–6 µm. Pileus less vividly coloured (more fulvous-grey brown), rarely bicoloured. Northern Europe (Fennoscandia) .......................................................... C. caesiophylloides 6* 7 7* 172 Spores strictly amygdaloid, densely verrucose, (8.5‒)9‒10 × (4.5‒)5–6 mm. Usually robust, with distinctly marginate bulb. Pileus often silvery-ibrillose from thin veil remnants. Honey smell weak or absent. Associated with Picea and Abies, mainly in S Fennoscandia and C Europe ........................................................................................... C. talimultiformis Spores (broadly) ellipsoid-subamygdaloid, with somewhat coarser (conluent-rounded), diffuse ornaments. Robust or not ............................................................................................. 8 8 8* 9 9* Pileus strongly apricot-redbrown, with distinct pattern of dark, hygrophanous spots. Spores ellipsoid-subamygdaloid, 8–9.5 × (4.5–)5–5.5 mm, weakly verrucose to almost smooth. Subcutis (hypoderm) with some distinctly zebra-striped encrusted hyphae. Honey smell weak or absent ............................................................................................... C. rufoallutus Pileus ochraceous brown-fulvous brown, usually without a pattern of hygrophanous spots. Subcutis (almost) without zebra-striped encrusted hyphae ................................................. 9 Spores 8–9.5 × (4.5–)5–5.5(–6) mm, ellipsoid-subamygdaloid; basidiomes often slender and with an indistinct bulb, almost without veil remnants; honey smell weak or absent. Mainly in Northern Europe. Widespread. Associated with Picea .............................. C. multiformis Spores 9–10 × 5.5–6.5 µm, (broadly) ellipsoid; basidiomes robust, with distinct, white veil remnants at bulb (and pileus) margin; with a distinct honey smell. Recorded from Picea and Larix-Betula forests. Very rare (E Siberia, E France) ................................ C. armenicorius The C. multiformis–C. rufoallutus subclade Cortinarius multiformis Fr. Figs. 3, 4, 5 Epicr. Syst. Mycol.: 263 (1838). Type: Sweden, Cortin. Fl. Photogr., CFP 445, herb. S; neotype designated in LiimaTainEn et al. (2014). Syn.: C. lundellii (M. M. Moser) M. M. Moser, Kleine Kryptogamenlora Bd IIb/2: 286 (1967) Phlegmacium lundellii M. M. Moser, Die Gattung Phlegmacium: 349 (1961) C. subhygrophanicus (M. M. Moser) M. M. Moser, Kleine Kryptogamenlora Bd IIb/2: 286 (1967) Phlegmacium subhygrophanicus M. M. Moser, Die Gattung Phlegmacium: 349 (1961) ?C. coniferarum (M. M. Moser) Moënne-Locc. & Reumaux, Atlas des Cortinaires III: Fiche 114 (1991) Phlegmacium multiforme (Fr.) Wünsche var. coniferarum M. M. Moser, Die Gattung Phlegmacium, p. 349 (1961) ?C. herbarum Rob. Henry, Bull. Soc. Mycol. Fr. 67(3): 270–271 (1951) ?C. pseudorugulosus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires XVI: 1096 (2006). Misapplied name: C. allutus Fr. s. auct. Icones: Brandrud et al. (1990–2014): A 85 (CFP 445; neotype), Bidaud et al. (2006): Pl. 579 (sub nom. C. lundellii), MoSEr (1961): Pl. IV (sub nom. C. lundellii, C. subhygrophanicus), Pl. I (sub nom. C. allutus). kriEGlSTEinEr & GmindEr (2010): p. 141 (sub nom. C. multiformis var. coniferarum). Pileus: (3‒)4–8 cm, (hemi-)spherical, then plano-convex, viscid, often radially rugulose with age; often with a dense pattern of hygrophanous spots (centre) or streaks/veins (near margin) (pseudohygrophanous structure), with age sometimes entire marginal zone hygrophanous, centre sometimes silvery-micaceous from thin veil remnants when young; rather uniformly (dark) ochraceous yellow, fulvous to almost red brown, often becoming pale ochraceous yellow with age. Lamellae: crowded, 4–8 mm broad, initially greyish white. Stipe: 5–12(–14) × 0.8–1.5 cm, usually slender, with a faintly to rather distinctly marginatebulbous base (®2.5 cm), often becoming quite non-marginate (almost cylindric) with age; glossy, white, turning distinctly brass-brown with age. Universal veil near the bulb very sparse, whitish at bulb margin. Context: white, somewhat (ochre)greyish hygrophanous-spotted (marbled) towards surface of stipe (apex). Smell none or faint of honey, at least in bulb when bruised. Macrochemical reactions: negative or indistinct. 173 Spores: 8–9.5 × (4.5–)5–5.5(–6) mm (MV = 8.8 × 5.3 mm), Q = 1.67, ellipsoid to subamygdaloid, pale, weakly to distinctly and rather densely to diffusely verrucose, warts irregular to slightly conluent, rounded, sometimes warts very lat and hardly visible in the outline of the spores. Photo: Geert Schmidt-Stohn Fig. 3 – Spore morphology of species in the C. multiformis subclade. 174 Spores 2000 : 1. Photo: Geert Schmidt-Stohn Fig. 4 – Cortinarius multiformis (Sweden, SSt10-068; +DNA) Scatter diagram: Geert Schmidt-Stohn Fig. 5 – Variation in spore size amongst species in the C. multiformis subclade. Each symbol = MV of spores measured per basidiome. 175 Pileipellis: duplex, at surface of thin, erect-entangled gelatinous, ±hyaline hyphae. Subcutis/ hypodermium subcellular, hyphae tightly cemented, with yellow brown parietal-encrusting pigment, forming an amber-like embedment. Thick, brown walls especially pronounced in corners between the cells; forming irregular-triangular «lenses», to more diffuse, brown pigments lumps. More narrow, longer transition hyphae between subcutis with and epicutis with yellow walls, and sometimes with zebra-striped encrustration. DNA (ITS) sequence: Fifteen of the European ITS sequences available, including the neotype, are identical. However, 1 collection shows a minor difference without support. Two North American (Canadian) collections of C. multiformis differ by 1 nucleotide from the European ones (data not shown). Habitat: Boreal (-boreonemoral) and (more rarely) montane-subalpine coniferous forests. Associated with Picea abies, mainly in oligotrophic-acid, mossy sites, often in large numbers in young plantations. Distribution: Frequent and widely distributed in the Nordic boreal spruce forest region (JEppESEn et al. 2012), including Estonia. Apparently more rare in the montane-subalpine regions of C Europe, but distribution poorly known due to confusion with C. talimultiformis. So far material of C. multiformis s. str. outside Northern Europe documented with ITS sequences only from the Prealps (N Italy, Austria) and one collection from Spain (see below). Probably also recorded from France; the plates of C. lundellii and C. herbarum, possibly also C. pseudorugulosus in Bidaud et al. (2006) apparently depict our species (material not investigated). Collections examined: NORWAY. Oppland: Øystre Slidre, Reieåsen, TEB 410-13, DB-NO-33 (herb. O) (DNA*). Øystre Slidre, Volbufjorden, TEB 413-13, DB-NO-37 (herb. O) (DNA*). Vestre Slidre, Mosåni, TEB 186-07 (herb. O) (DNA*). Gjøvik, Biri, K. H. Brandrud & TEB 102-85 (herb. O). Lunner, S. Oppdalen, TEB 205-77, 277-78 (herb. O). Jevnaker, Sinnerseter, E. Bendiksen & TEB 40-81 (herb. O). Hordaland, Voss, T. Niskanen et al., TN05-247 (herb. H) GenBank No. KF732626 (DNA*). SWEDEN. Ångermanland: Häggdånger, Cortinarius, Flora Photographica, CFP 445 (neotype, herb. S) (DNA*), CFP 88 (to herb. S). Halland: Mästocka, 9. July 1998, T. Münzmay H.15.98 (herb. TUB 4848) (DNA). Dalarna: Mora, T. Münzmay 11 Sept. 2007 (herb. TUB 9481) (DNA). Revnes, K. Soop 18. Sept. 2010, KS-CO1974 (herb. S F189954) (DNA*). Jämtland: Ragunda, TEB 15-93 (herb. O). Ragunda, Bispgården, A. Taylor, AT2004187 (herb. UPS), UNITE No. UDB002163 (DNA*). Offerdal, TEB 25-82, 26-82 (herb. O). Medelpad: Borgsjö, CFP 16 (to herb. S), K. Leonardsson & TEB 47-82 (herb. O). Borgsjö, Jämtgaveln, G. Schmidt-Stohn, SSt10-068 (DNA*). Svartede W of Sundsvall, TEB 66-82 (herb. O). Norrbotten: Gällivare, K. Høiland & TEB 131-81 (herb. O). ITALY. South Tyrol: Sarntal, Reinswald, F. Bellù 21. Sept. 2011, Bellu21-09-2011 (herb. BOZ) (DNA*). Corti, Grünbachalm, G. Turrini, TG1999-231 (herb. BOZ) (DNA*). Valdaora di mezzo/Mitterberg, G. Turrini, TG2001-098 (herb. Turrini) (DNA). Pochi di Salorno, Val Fredda, G. Turrini, TG1998-096 (herb. Trento) (DNA*). AUSTRIA. Gepatsch Stausee, SSt 09-017, G. Schmidt-Stohn (herb. TUB) (DNA*), Zillertal, M. Dondl, MD12-2012 (DNA*). SPAIN. J. Ballarà, JB 6591-09 (DNA). Comments: Cortinarius multiformis is characterized by its vivid and uniformly yellow brown pileus colour, resembling that of ripe Rubus chamaemorus berries. Furthermore the stipe is initially shiny white, slender and usually with only a poorly developed marginate bulb, often sitting in deep moss. The description and plate of C. lundellii in MoSEr (1961) well illustrates this often slender habit (see also Figure 4). Until recently, C. multiformis has usually been treated collectively, including C. talimultiformis, and possibly also C. armenicorius, C. frondosomultiformis and C. rufoallutus. These have overlapping morphological variation, but are quite well distinguished phylogenetically. Cortinarius talimultiformis is normally more robust, with a more distinctly and persistently marginate bulb, and on average possess a more vividly orange/red brown pileus and more whitish veil remnants on the pileus. Cortinarius armenicorius is paler, with more veil rem176 nants, and a distinct honey-like smell. The often co-occurring Cortinarius rufoallutus is more red brown, has a coarser, more pronounced hygrophanous spotted pileus surface, and early becomes brown spotted at stipe surface and in context. Pale forms of C. multiformis can sometimes be dificult to separate from the deciduous forest species C. talus and C. frondosomultiformis but these have a silky-micaceous-innately ibrillose structure of young pilei, and have a stronger smell of honey. Cortinarius talus has distinctly paler colours than C. multiformis when young and fresh, but C. frondosomultiformis and C. multiformis apparently might have overlapping colours, but is distinguished on the habitat (Quercus/Fagus versus Picea association). The European material available shows a minor variation in ITS sequence. Cortinarius frondosomultiformis comes out as sister species in the phylogramm, with 9–10 differences in ITS from C. multiformis (Figure 1). Another very close genotype («C. sp.» in Figure 1) is distinguished from C. multiformis by 5–6 nucleotide differences. Due to limited knowledge, this is preliminary regarded as cryptic species, without a formal, taxonomic recognition. Cortinarius frondosomultiformis Bellù, Brandrud & Dima sp. nov. Figs. 3, 5-8 MycoBank No. MB809862 Etymology: from latin frondosis and multiformis, meaning „a C. multiformis in frondose forests” Typus: Italy, South Tyrol, Kaltern east, leg. G. Turrini, TG 2000-218 (herb. BOZ, holotype) GenBank No. KM504516. Pileus: 5–7 cm, (hemi-)spherical, then plano-convex, viscid, initially slightly marbled-pseudohygrophanous of a dense pattern of small, darker spots, sometimes larger hygrophanous spots/ streaks near margin, margin often undulate and with a narrow, hygrophanous zone with age; sometimes slightly innately ibrillose towards margin when young; ochraceous yellow(-brown), somewhat paler towards margin and with age. Universal veil remnants sparse, but when young often with thin, silvery-micaceous-frosty remnants, especially at centre. Lamellae: crowded, with entire to later often sinuate-dentate margin, initially greyish white. Stipe: 4–7(–9) × 1–1.5(‒2) cm, robust to sometimes slender, with an usually distinctly marginate, rather broad bulb (®3 cm), silky-ibrillose to glossy, initially pure white, turning slightly brass-brown with age, especially when bruised. Universal veil at the bulb very sparse, white. Context: white, somewhat (ochre)greyish hygrophanous-spotted (marbled) towards surface of stipe (apex). Smell distinct of honey in bulb context. Spores: 7‒8.5(‒9) × 4‒5(‒5,5) mm (MV = 7.9 × 4.6 mm), Q = 1.71, ellipsoid to subamygdaloid, pale, distinctly and rather densely to more coarsely, diffusely verrucose, warts rounded and often conluent. Pileipellis: duplex, cutis of gelatinous, loosely erect-entangled, hyaline hyphae at surface, subcutis/hypodermium subcellular, hyphae tightly cemented, with yellow brown to golden yellow parietal-encrusting pigment, forming an amber-like embedment. Very thick walls in corners between the cells forming brown, irregular-triangular “lenses” to more diffuse pigment lumps. DNA (ITS) sequence: All 3 ITS sequences generated in this study are identical. Habitat: Nemoral-colline to montane, thermophilous deciduous forests. Apparently associated mainly with Quercus spp., probably also with Fagus sylvatica, Carpinus betulus, Castanea sativa and possibly Abies alba, in richer, but not typically calcareous sites. Distribution: Hitherto known only from a few collections from N Italy (South Tyrol; three collections, two sites). The absence from Western Europe and temperate parts of Northern Europe is remarkable: many specimens from this group have been ITS investigated from these regions, with no match on this one so far. 177 Photo: Gianni Turrini Fig. 6 – Cortinarius frondosomultiformis Holotype (N Italy, TG2000-218; +DNA) Photo: Gianni Turrini Fig. 7 – Cortinarius frondosomultiformis Holotype (N Italy, TG2000-218; +DNA) 178 Photo: Tobias G. Frøslev Fig. 8 –Cortinarius frondosomultiformis (N Italy, TF 2000-086; +DNA) (The mossy background is not from the collecting site.) Collections examined: ITALY. South Tyrol, Kaltern east, Montigglerwald near Eppan (ca. 450–470 m a.s.l.), leg. G. Turrini 22. Oct. 2000, TG 2000-218 (herb. BOZ, holotype) (DNA*); Kaltern west, Matschatsch (route to Mendelpass), 882 m a.s.l., T. Frøslev & T. S. Jeppesen, TF 2000-086 (herb. C) (DNA*); Matschatsch, leg. A. Cester, det. F. Bellù (as C. talus) 07.11.2011 (herb. BOZ 2011/71) (DNA*). Comments: Cortinarius frondosomultiformis is characterized by its (i) small spores, (ii) moderately pale ochraceous yellow pileus, (iii) frosty-micaceous appearance of pileus centre, (iv) robust specimens with a rather distinctly marginate bulb and (v) a distinct honey-like smell. It is furthermore the only species in sect. Multiformes which seems restricted to thermophilous deciduous forests (with Quercus-Carpinus-Fagus), possibly including Abies alba forests, which often can harbour frondose forest species. All three collections examined have smaller spores than related taxa, and if constant, this is an important differential character. The species is macroscopically intermediate between C. multiformis/C. talimultiformis on the one hand and C. talus on the other; the pileus is darker than that of C. talus and paler than that of C. multiformis and C. talimultiformis. The frosty-micaceous-marbled surface structure is more pronounced than in C. multiformis and C. talimultiformis but less than in C. talus. However, the pileus colour (and the parietal pigment of subcutis) when young comes closer to C. multiformis than C. talus, whereas the distinct smell (honey) and the habitat (Quercus-Fagus forests) comes closer to C. talus. Phylogenetically, C. frondosomultiformis is most similar to C. multiformis, with 9–10 nucleotide differences. 179 Cortinarius talimultiformis Kytöv., Liimat., Niskanen, A.F.S. Taylor & Sesli Figs. 3, 5, 9 Persoonia: 33 (2014). Type: Sweden, A. Taylor, AT2004096, herb. UPS, holotype; S, isotype Syn.: Cortinarius aurantionapus Bidaud & Reumaux var. similis Moënne-Locc., Atlas des Cortinaires XVI: 1096 (2006). Misapplied name: C. multiformis Fr. s. auct. Icones: Bidaud et al. (2006): Pl. 596 (holotype), 597 (both sub nom. C. aurantionapus var. similis). Pileus: 4–10 cm, (hemi-)spherical, then plano-convex, viscid; uniformly fulvous brown to more orange brown, and slightly more red brown at centre. Universal veil sparse, but when young often whitish ibrillose towards margin, and sometimes also silvery-micaceous at centre. Lamellae: crowded, initially greyish white. Photo: Bálint Dima Fig. 9 –Cortinarius talimultiformis (Hungary, DB2831; +DNA) Stipe: 4–6(–10) × 1–2 cm, usually robust, with a more or less distinctly marginate bulb (®3.5 cm), glossy-silky, white, turning slightly brass brown with age. Universal veil at the bulb sparse, sometimes more distinct, white. Context: white, somewhat (ochre)greyish hygrophanous-spotted (marbled) towards surface of stipe (apex), sometimes brownish spotted in the bulb. Smell indistinct or faint of honey in the bulb when cut. Spores: (8.5‒)9‒10 × (4.5‒)5–6 mm (MV = 9.1 × 5.4 mm), Q = 1.68, (sub)amygdaloid, sometimes acutely amygdaloid, sometimes with a slight suprahilar depression, distinctly and rather densely verrucose, warts rounded and sometimes conluent. Pileipellis: duplex, cutis of gelatinous, loosely erect-entangled, hyaline hyphae at surface. Subcutis/hypodermium subcellular, hyphae tightly cemented, with strong yellow brown parietal180 encrusting pigment, forming an amber-like embedment, often with thick, brown irregular-triangular «lenses» between the cells, grading into more diffuse, brown pigment-lumps. DNA (ITS) sequence: All 11 ITS sequences included in our analysis (5 generated in this study) are identical. Although with a somewhat lower bootstrap value in our RAxML analysis (Figure 1), it is still well-separated phylogenetically from the other species in sect. Multiformes. The study of LiimaTainEn et al. (2014) also clearly supports the separation of this species. Habitat: Montane-subalpine and (more rarely) boreal (-boreonemoral) and nemoral coniferous forests. Associated with Picea and Abies, in oligotrophic to richer/calcareous sites, including young plantations. Very rarely also in mixed broad-leaf forests (with Picea/Abies?). Distribution: Widely distributed and probably not infrequent in Central Europe, including The Black Forest region, The Prealps, the Jura and W Hungary (cfr. Bidaud et al. 2006, as C. aurantionapus var. similis). Recorded in S Europe southeast to Turkey (LiimaTainEn et al. 2014). Apparently rarer in N Europe (collections conirmed by ITS sequences only one from Norway, one from Sweden but several from Finland; LiimaTainEn et al. 2014). Frequency and distribution uncertain due to confusion with the closely related C. multiformis s. str. and C. rufoallutus. Collections examined: GERMANY. Baden-Württemberg: Titisee-Neustadt, 9. Sept. 2002, G. Saar (herb. TUB 011864, GenBank No. AY669532 as «C. allutus») (DNA*); Schwaben, Ehingen a.d. Donau, 28. Sept. 2010, G. Schmidt-Stohn, SSt 10-188 (herb. TUB) (DNA*). Niedersachsen: Ahlsburg, Dörrigsen, G. Schmidt-Stohn, SSt 09-040 (herb. TUB) (DNA*). Bavaria: Bad Hindelang, Oberjoch, 25. Sept. 2007, G. Saar, SG 01722 (herb. TUB 019722) (DNA); Oberjoch, 2. Oct. 2002, G. Saar, SG 0677 (herb. TUB) (DNA). ITALY. Trentino: Tovare Malè, val di Sole, 29. Aug. 2002, C. Rossi, det. F. Bellù (as C. allutus) (herb. Rossi29-08-2002; photo no. 5897) (DNA*). South Tyrol: Pochi di Salorno, Val Fredda, 19. July 1998, G. Turrini, TG 1998-096 (herb. Turrini, as C. allutus; photo no. 754) (DNA). Brunico, Amaten, Montassilone, Taurerntal, 10. Aug. 1999, G. Turrini, TG 1999-246 (herb. Turrini, as C. multiformis; photo no. 1556) (DNA*). HUNGARY. Vas, Felsőszölnök, L. Albert, B. Dima, DB2831 (herb. BP) (DNA*). SWEDEN. Uppland: Hässelby near Uppsala, Andy Taylor, AT2004036 (herb. UPS, holotype; S isotype) (DNA*). FINLAND. Kainuu: Puolanka, T. Niskanen & K. Liimatainen, TN01057 (herb. H) (DNA). Pohjois-Häme, Virrat, Killinkoski, 30. Aug. 1966, H6032747 (herb. H), GenBank No. KF732584 (DNA*). Comments: Cortinarius talimultiformis is characterized by its comparatively large, rather strictly amygdaloid spores and vivid and uniformly fulvous-orange brown pileus with ine whitish-silvery ibrillose veil remnants when young. Furthermore, the stipe is initially shiny white, short and robust and usually with a distinctly marginate bulb. Until recently, C. multiformis has usually been treated collectively, including C. talimultiformis and C. multiformis s. str., which now are documented to be well distinguished phylogenetically. However, these are dificult and sometimes probably impossible to distinguish morphologically. The somewhat larger and strictly amygdaloid spores of C. talimultiformis (versus more subamygdaloid-ellipsoid in C. multiformis) might be the best differential character, but size and probably also shape of the spores can be overlapping. Also the robust habit, including a rather wide, more or less distinctly marginate bulb and the initially white veil remnants on pileus of C. talimultiformis will often be distinguishing towards C. multiformis. Most of these macro-features are well demonstrated in the plates 596, 597 and front cover in Atlas des Cortinaires XVI (Bidaud et al. 2006; as C. aurantionapus var. similis) and in our photo of sequenced material (Figure 9). Cortinarius talimultiformis occur sometimes in Picea plantations in nemoral regions (cfr. Figure 9), and might in such areas possibly occur also in deciduous tree-dominated forests, and our species can then be dificult to distinguish from C. frondosomultiformis. The latter, however has paler, not so vivid orange-brown pileus colours, and smaller spores. 181 Cortinarius talimultiformis can also be dificult to distinguish from C. rufoallutus, but the latter has smaller spores, a hygrophanous-spotted pileus, with zebra-striped encrusted pigment in pileipellis and pigment that often also goes into the pileus trama, and the latter also becomes more strongly brass brown spotted on stipe and in the context. Cortinarius talimultiformis is most closely related to C. multiformis, distinguished by 6 nucleotide differences in the ITS regions. Cortinarius rufoallutus Rob. Henry ex Bidaud & Reumaux Figs. 3, 5, 10, 11 Atlas des Cortinaires XVII: 1095 (2006). Type: France, leg. P. Moënne-Loccoz & J. Melot, PML 635, herb. PC, holotype. Syn.: C. rufoallutus Rob. Henry nom. inval., Bull Soc. Mycol. Fr. 74(3): 329–331 (1958) C. allutus var. rufescens Rob. Henry nom. inval., Bull Soc. Mycol. Fr. 55(1): 66–68 (1939). Misapplied name: C. aurantiacus M. M. Moser s. auct. Icones: Bidaud et al. (2006): Pl. 580 (holotype), Soop (2011): Fig. 22. Pileus: 5–8(‒10) cm, (hemi-)spherical, then plano-convex, viscid-glutinous; usually with distinct, dense, dark hygrophanous spots and sometimes more elongated streaks near margin, centre sometimes silvery-micaceous from thin veil remnants when young; rather uniformly dark and vividly fulvous brown, apricot to almost red brown. Lamellae: crowded, 4–8 mm broad, initially greyish white. Stipe: 5–7(–9) × 1–2 cm, with a usually distinct marginate-bulbous base (®3.5 cm); rather robust/thick-leshed (stipe length/width ratio 3.5–5.5 cm); ibrillose, whitish, but soon turning turning strongly, spot-wise brass brown, especially when bruised, entire stipe (dark) brown with age. Universal veil near the bulb very sparse, initially whitish at bulb margin. Context: whitish, with with a distinct thin, brown zone beneath pileipellis and in stipe cortex, also somewhat ochraceous grey hygrophanous-spotted (marbled) towards surface of stipe (apex). Smell none or faint of honey in bulb when bruised. Photo: Karl Soop Fig. 10 – Cortinarius rufoallutus (Sweden, KS CO246; +DNA) 182 Photo: Tor Erik Brandrud Fig. 11 – Cortinarius rufoallutus (Norway, TEB 460-11; +DNA) Spores: 8–9.5 × (4.5–)5–5.5 mm (MV = 8.5 × 5.1 mm), Q = 1.68, ellipsoid to subamygdaloid, pale, weakly to distinctly and rather densely to diffusely verrucose, warts irregular to slightly conluent, rounded. Pileipellis: duplex, at surface of thin, erect-entangled gelatinous, ±hyaline hyphae. Subcutis/ hypodermium subcellular, hyphae tightly cemented, with strong, yellow brown parietal-encrusting pigment, forming an amber-like embedment. Very thick walls in corners appear like brown irregular-triangular «lenses», or as more diffuse, oleiferous, brown pigments lumps. Some more narrow, long hyphae (4-8 um wide) also with distinct, zebra-striped, encrusted pigment (especially at surface; transition to epicutis). DNA (ITS) sequence: All European ITS sequences available (5), including the holotype, are identical. Habitat: Boreal (-boreonemoral) and montane-subalpine coniferous forests. Associated with Picea abies, mainly in oligotrophic-acid, mossy sites, sometimes in young plantations. Distribution: Rare but widespread in the Nordic boreal spruce forest region (pers. obs., cfr. Soop 2011). Apparently very rare in the montane-subalpine regions of C Europe but sequenced material is documented from France (holotype) and from Italy. Distribution poorly known due to confusion with C. multiformis and C. talimultiformis. Collections examined: NORWAY. Oppland: Øystre Slidre, Volbufjorden, TEB 415-13, DBNO-38 (herb. O) (DNA*). Øystre Slidre, Heggen church, TEB 460-11 (herb. O) (DNA*). Øyer, Skardsmoen, M. K. Brandrud & TEB 217-11 (herb. O). Lunner, S. Oppdalen, TEB 1-89 (herb. O). Gran, Øytjernet, E. Bendiksen & TEB 74-80 (herb. O), Buhammeren, TEB 115-79 (herb. O). Gjøvik, Glæstadtj., K. H. Brandrud & TEB 107-85 (herb. O). Akershus: Nannestad, Hornsjøen, TEB 211-78 (herb. O). Nordmokorset, K. H. Brandrud & TEB 8-86 (herb. O). Oslo: Grorud, TEB 688-11 (herb. O). 183 SWEDEN. Dalarna, K. Soop KS-CO246 (herb. S) (DNA), KS-CO747 (herb. S) (DNA). ITALY. South Tyrol: Anterselva, leg. C. Rossi 02. Aug. 2006 (herb. Rossi02-08-2006, as C. multiformis) (DNA*). Comments: Cortinarius rufoallutus is macroscopically fairly characteristic with its robust basidiomes, warm, vivid fulvous-orange brown-red brown pileus, usually with distinct, small hygrophanous spots, and a stipe that turns vividly brass brown. A vivid ochre brown line beneath pileipellis is also characteristic. The species often co-occur with C. multiformis, but is much rarer. The mentioned features normally distinguish it from C. multiformis when specimens are young and fresh, but sometimes their colours might overlap. However, the shape of stipe hardly overlap; being more slender in C. multiformis, with a length/with ratio of 5.0–9.5, versus 3.5–5.5 in C. rufoallutus. Furthermore, C. rufoallutus can be separated also microscopically by presence of distinct zebra-striped encrusted hyphae in the transition between epicutis and subcutis. This feature also distinguishes it from C. talimultiformis, which also might be vividly apricot-red-brown, but rarely with a hygrophanous-dotted structure. The latter also often have more veil remnants on pileus. Phylogenetically, C. rufoallutus comes closest to C. talimultiformis and C. multiformis, but is well-distinguished from these by 18-20 nucleotide differences. After introduced by HEnry (1939) as C. allutus var. rufescens nom. inval., this species has been little known and described. The senior author has since 1978 recognized this one and collected it as C. multiformis var. rufescens, but has doubted speciic rank, due to more or less continuous morphological variation with C. multiformis s. str. However, most of the taxa treated as varieties by Scandinavian authors in the 1980–1990ies (cfr. Brandrud et al. 1990–2014) have later showed to be well-supported species phylogenetically. Nevertheless, the taxon was treated as a separate species in early versions of «Cortinarius in Sweden» by K. Soop under the name C. aurantiacus, later changed to C. rufoallutus (Soop 2011). The plate illustrates well the brownspotted basidiomes of our species. The name C. rufoallutus, introduced by HEnry (1958), was validated by Bidaud et al. (2006) and the type material and plate correspond well morphologically and phylogenetically with our concept. Cortinarius melleicarneus Kytöv., Liimat., Niskanen & Brandrud Fig. 3 Persoonia: 33 (2014). Type: Estonia. Hiiumaa: Sarve, Soonlepa, 16 Sep 2001, I. Kytövuori 01-053, herb. H, holotype Icone: LiimaTainEn et al., Persoonia 33 (2014) (holotype). Pileus: 4–10 cm broad, hemispherical to convex, with rather persistently incurved margins, then expanded sometimes somewhat silvery-silky from ine veil remants, cream-coloured, pale yellow brown, honey brown to more grey brown or lesh-coloured, with hygrophanous streaks or patches/zone towards margin. Hygrophanous zone somewhat darker grey brown, almost with an olivaceous brown tinge. Lamellae: crowded, irst pale greyish white, later pale greyish brown. Stipe: short/robust; 5–7(–9) cm long, 1.2–2 cm thick at apex, 2–3 cm wide at base, clavate or with a somewhat marginate bulb, white. Universal veil: very sparse at bulb margin, white. Context: in pileus pale yellow brown, marbled hygrophanous lesh-coloured, in stipe white. Smell: not recorded. Spores: (7.5‒)8–8.5(‒9) × (4‒)4.5–5 µm, MV = 8.3 × 4.6 µm, Q = 1.81, narrowly (sub)amygdaloid, sometimes with a faint suprahilar depression, medium (to fairly strongly), densely to fairly coarsely verrucose, warts rounded and more or less conluent. Pileipellis: duplex, at surface of thin, erect-entangled gelatinous, ±hyaline hyphae. Subcutis/ hypoderm subcellular, hyphae tightly cemented, with pale yellowish parietal, amber-like pigment, a few (more narrow) hyphae with zebra-striped encrusted pigment occur intermixed. 184 DNA/ITS sequence: Sequences from the two, known collections are identical (cfr. LiimaTainEn et al. 2014). Ecology: With thermophilous deciduous trees (Corylus, Quercus) on calcareous ground, including near-shore sandy shell-beds (remnants of sandy soil on both collections). Distribution: Known from the boreonemoral zone of S Scandinavia-Balticum (Estonia and Norway). Collection examined: ESTONIA. Hiiumaa: Sarve, 16. Sept. 2001, I. Kytövuori, IK01-053 (herb. H, holotype), GenBank No. KF732577 (DNA*). NORWAY. Aust-Agder, Grimstad, Fevik, 21. Sept. 1994, I-L. Fonneland, det. T. E. Brandrud 86-94, O 125960 (herb. O; sub nom. C. arenisilvae), GenBank No. AY669533 (DNA*). Comments: Cortinarius melleicarneus is characterized by robust, short-stiped basidiomes, honey-brown pileus with initially pale centre, and slightly smaller and narrower spores than related taxa. It is further characterized by habitat in broad-leaf forests on calcareous ground. Cortinarius talus resembles in pale pileus colours and habitat, but this usually has an innately ibrillose pileus structure, and rarely possess short-stemmed, compact basidiomes. No honey-smell was noted in the two collections, and the lack of this smell might be a diagnostic character towards the usually strongly honey-smelling C. talus. In dry conditions, this might resemble the more ibrillose C. areni-silvae (Brandrud) Brandrud which also occur in sandy soils, and the Norwegian collection was originally determined as C. areni-silvae (based on dry material and collectors notes). ITS sequences of the two C. melleicarneus collections are identical. It differs by 11 nucleotides from C. talimultiformis, 14 from C. talus, and 15 from C. multiformis. The C. talus subclade Cortinarius talus Fr. Figs. 12, 13-15 Epicr. Syst. Mycol.: 263 (1838). Type: Sweden, Cortin. Fl. Photogr. CFP 832, herb. S; neotype designated in LiimaTainEn et al. (2014). Syn.: C. ochropallidus Rob. Henry, Bull. Soc. Mycol. Fr. 52(2): 153 (1936) C. melliolens Jul. Schäffer ex P. D. Orton, Trans. Brit. Mycol. Soc. 43(2): 2010 (1960) C. ochropudorinus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires XVI: 1097 (2006) C. crenulatus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires XVI: 1097 (2006) C. pseudotalus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires XVI: 1098 (2006) C. aurantionapus Bidaud & Reumaux, Atlas des Cortinaires XVI: 1096 (2006) C. pudorinus Reumaux, Atlas des Cortinaires XVI: 1098 (2006) C. pseudominor Rob. Henry ex Reumaux, Atlas des Cortinaires XVI: 1098 (2006). Misapplied name: C. multiformis Fr. sensu Moser. Icones: Brandrud et al. (1990–2014): B 47 (CFP 835; neotype). Bidaud et al. (2006): Pl. 601–601 (sub nom. C. ochropudorinus) 603, 604, 605, 608 (sub nom. C. pseudotalus, C. gregis), 609 (sub nom. C. minor, C. pseudominor), 610 (sub nom. C. leptocephalus), 611 (sub nom. C. minor), 612 (sub nom. C. ochropallidus). Pileus: 3–9 cm broad, hemispherical to plano-convex, viscid, centre often silvery-«frosty» when young due to a thin cover of veil remnants, towards margin initially often very inely innately ibrillose-marbled or with hygrophanous spots or streaks; ochraceous yellow, sometimes almost whitish when young, sometimes almost ochraceous brown when young (if the whitish supericial hyphae are washed away). Lamellae: crowded, pale greyish white when young. Stipe: 4–8(–10) x 0.8–1.5 cm, with a (slightly) marginate bulb (→ 2.5 cm), glossy, whitish, then brass-brown spotted. Universal veil: white, sparse. Context: whitish. Smell: distinctly to strongly of honey when cut. 185 Photo: Geert Schmidt-Stohn Fig. 12 – Spore morphology of species in the C. talus and C. pallidirimosus-C. caesiolamellatus subclades. Spores 2250:1. 186 Spores: (7–)7.5–9.5 × 4.5–5.5 µm, (MV = 8.25 × 4.95 µm, Q = 1.68), amygdaloid to ellipsoid, sometimes broadly amygdaloid-ellipsoid, pale, weakly to moderately and rather densely verrucose, warts rounded and often somewhat conluent-diffuse. Pileipellis: duplex, at surface of thin, erect-entangled gelatinous, ±hyaline hyphae. Subcutis/ hypoderm subcellular, hyphae tightly cemented, with (very) pale yellowish parietal, amber-like pigment. DNA/ITS sequence: all 11 available sequences from N and S Europe are identical, while one North American collection differs by 2 nucleotides in the ITS regions (Figure 1). Habitat: in nemoral to montane/subalpine deciduous forests, sometimes also low alpine-subarctic heaths. In northern and subalpine regions of Scandinavia mainly associated with Betula pubescens, sometimes with Betula nana; elsewhere associated with Fagus, Quercus, Carpinus, Corylus and Betula pendula, in rich to oligotrophic forests (very rarely in calcareous sites together with calciphilous Phlegmacium species). Distribution: Occasional to rather frequent and widespread in Europe; from northern Scandinavia to the Mediterranean Sea. Also known from Greenland (KnudSEn & BorGEn 1987; with Betula pubescens) and from W North America (sequenced material, cfr. Figure 1). Probably with a wide, circumpolar distribution. Photo: Tor Erik Brandrud Fig. 13 – Cortinarius talus (Norway; from northern Betula pubescens forest; TEB 184-78) Collections examined: NORWAY. Oppland: Vågå, Gjende, TEB 184-78, 105-80 (herb. O). Hedmark: Åmot, Osdammen, R. Halvorsen & E. Bendiksen 528-80 (herb. O). Buskerud: Hol, Vikastølen, TEB 174-79 (herb. O). Vestfold: Tønsberg, Slagentangen, TEB 86-82 (herb. O). Østfold: Råde, Kajalunden, Ø. Weholt & TEB 610-80 (herb. O). Hvaler, Vesterøya, TEB 440-81 (herb. O). SWEDEN. Jämtland: Ragunda, Cortinarius Flora Photographica, CFP 832 (TEB 21-93) (neotype, herb. S) (DNA*). Undersåker, Välliste, TEB 85-83 (herb. O). Medelpad: Borgsjö, SSt 187 10-033 (DNA*), TEB 139-10 (DNA*), TEB 140-10 (SSt 10-038) (DNA*). Dalarna: Gesunda, near Mora, 11. Sept. 2007, G. Saar, T. E. Brandrud (herb. TUB) (DNA). Ångermanland: CFP 81, CFP 322 (to herb. S). Norrbotten: Gällivare, K. Høiland & TEB 159-81 (herb. O). Kiruna, K. Høiland & TEB 191-81 (herb. O). Småland: Femsjö, M. Moser & TEB 140-79 (herb. O). Scania: Degerberga, TEB 99-88 (herb. O). Ivö, TEB 312-87 (herb. O). DENMARK. Sealand: TF 1999-035 (herb. C) (DNA*). GERMANY. Baden-Württemberg: Tübingen, 16. Oct. 2001, G. Saar, SG 0613 (herb. TUB 19713) (DNA). Saarland: Schmelz, 30. Sept. 2001, T. Münzmay, TM H.32.01 (herb. TUB 011877) (DNA). Thüringen: Schellroda, 12. Oct. 2009, leg. Felix Hampe FHC010 (DNA). AUSTRIA. Burgenland: St. Margarethen, 2. Oct. 2003, G. Saar (herb. TUB 19737) (DNA). FRANCE. Rhone-Alps: Ain, Ordonnaz, TEB 51-91 (herb. O). Photo: Geert Schmidt-Stohn Fig. 14 – Cortinarius talus (Sweden; from northern Betula pubescens forest; SSt10-033; specimens with dark pileus, illustrating the large, infraspeciic variation; +DNA) Comments: Cortinarius talus is characterized by the pale, marbled-silvery-frosty-innately ibrillose pileus, usually strongly honey-like smell and a stipe with rather distinct marginate bulb. The marbled-frosty appearance of pileus centre when young resembles that of Cortinarius (Rozites) caperatus, the innately ibrillose structure towards margin reminds of e.g. C. riederi. The species is distinguished from C. multiformis by the paler, innately ibrillose pileus, the stronger honey smell and the habitat in deciduous forest. Cortinarius frondosomultiformis, also occurring in (thermophilous) deciduous forests might be more dificult to distinguish; it often possess a marbled-frosty pileus centre, but rarely an innately ibrillose margin. Furthermore, the colour beneath the pale, «frosty» surface hyphae is initially darker yellow brown in C. frondosomultiformis than in C. talus. Cortinarius talus is further distinguished from the northern C. pallidirimosus by the more robust basidiomes with a more marginate bulb. The C. talus populations of northern Betula 188 pubescens forests have also distinctly smaller and more broadly ellipsoid spores (7‒9 × 4.5‒5.5 µm) than those of C. pallidirimosus (9‒10 × 5.5‒6 µm) (see below). Scatter diagram: Geert Schmidt-Stohn Fig. 15 – Variation in spore size of C. talus. Each symbol = MV of spore measures per basidiome. Northern = middle/northern boreal (subalpine) Betula pubescens populations. Southern = more southern Fagus-Quercus-Carpinus-Corylus-Betula pendula populations. The spore size and shape of C. talus shows a larger variation than normally seen in sect. Multiformes. Whereas species such as C. multiformis and C. rufoallutus has spores with mean values ranging from approx. 8‒9.5 × 4.75‒5.75 µm, the C. talus mean values ranges from 7.25‒9.7 × 4.4‒5.4 µm (Figure 15). The populations of northern Fennoscandia, from northern/middle boreal Betula pubescens forests (or mixed birch-spruce forests) normally possess small and rather broad spores, with average values ranging from 7.25‒8.75 × 4.5‒5.25 µm (MV = 7.9 × 4.9 µm, Q = 1.63), whereas the populations of southern boreal-boreonemoral-nemoral-montane Betula pendula-Fagus-Quercus-Carpinus forests normally haver larger and more narrow spores with average values of 8–9.5 × 4.5–5.25 µm (MV = 8.6 × 5.0, Q = 1.73). These populations have identical ITS sequences, but this ecogeographically correlated spore variation might yet relect some population genetic differentiation, and these variants should be investigated by population genetic methods (e.g. using RAD sequencing, DavEy & BlaxTEr 2010). Furthermore, this spore size differentiation should be better documented by more samples and more measurements. For instance, it seems that some southern Fagus-Quercus associated C. talus populations included in Bidaud et al. (2006) possess small and wide spores, especially the C. ochropudorinus and C. pseudominor variants (with spores 6.5‒8 × (4‒)4.5‒5 µm). Conclusively, for the time being, we have decided not give these spore variants any taxonomic rank. Cortinarius talus can be rather variable also in macrocharacters such as size and in pileus col- 189 our; for instance in dry weather the whitish, supericial (veil) hyphae may be more pronounced than when exposed to moist weather (compare Figure 13 versus 14). These features certainly relect only a habitat-based, phenotypic variation, and are not correlated with genetics or geographical patters. Based on this variation, C. talus has been splitted in many species, but according to the morphological variation seen, as well as phylogenetic studies of a number of type specimens (see LiimaTainEn et al. 2014), we conclude that e.g. at least 9 of the species presented in Bidaud et al. (2006) are within the variation spectrum of our polymorphous C. talus. The following new taxa introduced in Bidaud et al. (2006) are furthermore identical in ITS sequences with C. talus, and is here regarded as synonyms: C. aurantionapus, C. ochropudorinus, C. crenulatus, C. pudorinus, C. pseudotalus and C. pseudominor (cfr. Figure 1). The C. pallidirimosus–C. caesiolamellatus subclade Cortinarius pallidirimosus Kytöv., Liimat. & Niskanen Fig. 12 Persoonia: 33 (2014). Type: Finland, I. Kytövuori 95-585, H6035694 (herb. H, holotype; NY, isotype. Icone: LiimaTainEn et al., Persoonia 33 (2014) (holotype). Pileus: 3–9 cm broad, hemispherical to plano-convex, viscid, very inely innately ibrillosemarbled or with hygrophanous streaks; whitish to cream-coloured, centre ochraceous yellow, becoming more ochraceus brown with age. Lamellae: crowded, pale greyish white when young, later pale brown. Stipe: 6–10(–13) × 0.7–1.5 cm, with a clavate to faintly marginate bulb (→ 2.5 cm), whitish, then very pale brownish. Universal veil: white, sparse. Context: white. Smell: of honey when cut. Spores: 9–10(‒10.5) × 5–6 µm, (MV = 9.6 × 5.6 µm, Q = 1.72), (sub)amygdaloid, rarely more ellipsoid, medium strongly and rather densely verrucose, warts rounded and sometimes coalescent. Pileipellis: Epicutis with a gelatinous layer of 1.5–3 µm wide, smooth, colourless hyphae. Hypoderm well-developed, subcellular, with very pale yellowish parietal, amber-like pigment, or almost colourless. DNA/ITS sequence: The C. pallidirimosus complex shows a comparative large sequence variation. Two sequences, both from Finland (including type material) are identical. Two sequences are slightly deviating (3 nucleotide differences from the type), whereas one sequence is more different (6 nucleotide differences (cfr. Figure 1). Habitat: Middle to northern boreal, in rich to oligotrophic mixed forests with Betula pubescens and Picea abies or Pinus sylvestris. Probably associated with Betula. Distribution: So far known from a number of localities in C/N Finland, a couple in C/N Sweden and one station from N Norway (LiimaTainEn et al. 2014). Collections examined: FINLAND. Kainuu: Puolanka, Väyrylä, Körölä, 15. Sept. 1997, I. Kytövuori, IK97-1523, H6035715 (herb. H) (DNA); Perä-Pohjanmaa: Tervola, Louepalo, 23. Aug. 2007, I. Kytövuori, IK07-692, H6000925 (herb. H) (DNA*). Inari Lappi: Utsjoki, Kevo, Tsieskula, I. Kytövuori, IK95-540, (herb. H) (DNA). Comments: The species is not seen fresh by the authors, and the description of macrocharacters is based on LiimaTainEn et al. (2014). Cortinarius pallidirimosus is characterized by the (very) pale, innately ibrillose pileus, an almost non-marginate bulb and comparatively large spores. The slender habit and pale pileus makes it look somewhat like a C. pinophilus, which, however, almost lacks a gelatinous pileus surface, turns yellowish and is not genetically related. With its pale, 190 marbled-innately ibrillose pileus, distinct honey-smell and habitat in (mixed) Betula pubescens forests, the species resembles C. talus. The latter, however is distinguished on (i) the more robust basidiomes with a more marginate bulb, a iner and often more distinct innately ibrillose-marbled structure of of pileus surface, and smaller spores. The C. talus populations found in northern Fennoscandian birch-spruce forests where C. pallidirimosus occur, have distinctly smaller and more ellipsoid spores (7‒9 × 4.5‒5.5 µm) than those of C. pallidirimosus (9‒10 × 5‒6 µm). When mean values per collection is considered, there is hardly any overlap in spore size variation between C. pallidirimosus and the northern Betula C. talus populations. Phylogenetically, C. armenicorius is the closest sister species to C. pallidirimosus. It has 8 nucleotide differences in the ITS regions from the type of C. pallidirimosus, but due to some internal, genetic variation in the latter, the barcoding gap between these two are not large. Cortinarius armenicorius differs from the latter in a more vividly ochre yellow (-apricot) pileus colour, a more robust habit and a distinctly marginate bulb with rather thick veil remnants. Cortinarius pallidirimosus is also rather closely related to C. caesiophylloides (distinguished by 17 nucleotide differences), but this is easily recognized on bluish tinges on the lamellae and at stipe apex. Cortinarius pallidirimosus shows a comparatively large ITS sequence variation; two collections have 3 ITS sequence differences, and one collection shows 6 sequence differences (Figure 1). The former appears to be (micro)morphologically and ecologically quite similar to the type of C. pallidirimosus, and this ITS variation is thus so far regarded as infraspeciic. The latter might represent a separate taxon, but is not treated further due to limited knowledge on morphology. Cortinarius pallidirimosus is apparently a northern species in Europe, so far found only in N/C Finland, N Sweden and N Norway. According to LiimaTainEn et al. (2014), it is associated with Betula (B. pubescens). However, all inds referred by these authors are from mixed forests including conifers, and an association with conifers cannot be ruled out. It seems anyhow that Betula is the primary associate. Some inds are e.g. from dry, mixed birch-pine forests, and Pinus is a very rare associate among the Multiformes, and it seems most likely that the species here is associated with Betula. Cortinarius armenicorius Soop & Brandrud, sp. nov. Figs. 12, 16 MycoBank No. MB809871 Etymology: from Latin armeniacum and corium, «apricot peel», due to the appearance and colour of the pileus. Typus: France, Jura, Prénovel, in Picea forest, 6 Oct. 2008, leg. K. Soop, KS-CO1865 (herb. S (F251131), holotype). GenBank No. KM504515. Pileus: 2.5–4.5 cm, (hemi-)spherical, then plano-convex, viscid, not hygrophanous, glabrous; rather uniformly warmly apricot yellow to ochraceous yellow, becoming somewhat paler ochraceous yellow with age. Universal veil remnants rather prominent, leaving white, ibrillose patches towards margin. Lamellae: L = 60–70, rather crowded, 3–5 mm broad, initially greyish white, soon pale brown. Stipe: 3–4.5 × 0.8–1 cm, rather robust, with a rather distinctly (to faintly) marginate bulb (®2 cm), dry; white without any shade of blue, remaining whitish with only weak brownish spots where bruised. Universal veil white, forming a sheath around the bulb, fairly copious; cortina white. Context: white, somewhat (ochre)greyish hygrophanous-spotted (marbled) towards surface of stipe (apex). Smell strong of honey, especially when cut. 191 Spores: 9–10 × 5.5–6.5 µm, MV = 9.5 × 6.1 µm, Q = 1.58, (broadly) ellipsoid to subamygdaloid, medium strongly to rather strongly and rather coarsely verrucose, warts rounded and conluent. Pileipellis: duplex, cutis thin, surface hyphae gelatinous, loosely erect-entangled, hyaline. Subcutis/hypodermium hyphae tightly cemented, with rather strong, yellow brown parietalencrusting pigment, forming an amber-like embedment. Very thick walls between the cells in corners form distinct, irregular-triangular, brown pigment «lenses», grading into more diffuse, intracellular pigment lumps. Long and narrow transition hyphae between subcutis and epicutis with yellow walls (no zebra-striped encrustration observed). Photo: Karl Soop Fig. 16 – Cortinarius armenicorius Holotype (E France, KS CO1865; +DNA) DNA (ITS) sequence: The two ITS sequences available have one nucleotide difference. Habitat and distribution: So far known from France (Jura) and Russia (Sakha republic, Siberia). In Siberia reported from Larix gmelinii forest with Betula platyphylla, in the Frence Jura from Picea abies forest. Collections examined: FRANCE: Jura, Prénovel, in Picea forest, 6. Oct. 2008, K. Soop, KSCO1865 (F251131) (herb. S, holotype) (DNA*). RUSSIA: Sakha republic, Khangalassky Ulus, Myachei-Sise Mountains, Larix gmelinii forest with Betula platyphylla, 8. Aug. 1999, U. Peintner IB19990590 (herb. IB), GenBank No. AF325586, UNITE No. UDB001073 (DNA*). Comments: Cortinarius armenicorius is characterized by its comparatively large, more or less ellipsoid spores, vivid ochre yellow-apricot yellow pileus colours, the comparatively thick veil remnants at bulb and pileus margin, and the distinct/strong honey smell. Phylogenetically, this taxon comes closest to C. pallidirimosus. The two collections of C. armenicorius possess only 8 192 nucleotide differences in ITS regions from the holotype of C. pallidirimosus, however, morphologically, these sister taxa are easily distinguished, C. pallidirimosus being more slender-clavate, and with a much paler, innately ibrillose pileus without white veil patches at margin. According to available material, C. armenicorius is morphologically much more similar to C. talimultiformis and C. multiformis, but seems to be (i) smaller, (ii) less (pseudo)hygrophanous, (iv) having a thicker veil, and (v) a stronger honey odour than those taxa. Furthermore, the spores of C. armenicorius are larger and more strongly ornamented than those of C. multiformis, and are more (broadly) ellipsoid than those of C. talimultiformis. With one collection from the very little surveyed Eastern Siberia, one might expect this to be a primarily northeastern Eurasian species. However, the ind from French Jura shows that this is distributed not only in the northern boreal taiga belt, but also in the coniferous forest belt of the Jura-Prealps. The collection from the French Jura indicates that this is a coniferous forest species with an association with Picea abies. The collection from the larch-dominated taiga forest of E Siberia indicate an association with Larix gmelinii. As association with Larix spp. is hitherto not documented for Multiformes species. However, also the East Asian Betula platyphylla was present at the Siberian site, and an association with northern-eastern Betula species cannot be ruled out. The sister taxon C. pallidirimosus is apparently mainly associated with northern Betula. Cortinarius caesiolamellatus (Bidaud) Kytöv., Liimat., Niskanen, Brandrud, Frøslev & A.F. Taylor Figs. 12, 17, 18 Type: France., P. Moënne-Loccoz 3. Oct. 1993, PML4905, herb. PC, holotype. Syn.: Cortinarius rufoallutus var. caesiolamellatus Bidaud, Atlas des Cortinaires XVI: 1095 (2006). Cortinarius multiformis var. caesiophyllus Moënne-Locc., Atlas des Cortinaires XVI: 1095 (2006). Type: France, P. Moënne-Loccoz 3. June 1988, PML882, herb. PC, holotype. Icones: Bidaud et al. (2006): pl. 581 (sub nom. C. rufoallutus v. caesiolamellatus), 587 (sub nom. C. multiformis var. caesiophyllus). Soop (2011): ig. 21 (sub nom. C. multiformis var. cyanoallutus ined.). ?kriEGlSTEinEr & GmindEr (2010): p. 81 (sub nom. C. allutus). Pileus: 4–8 cm, (hemi)spherical, then plano-convex, sometimes radially rugulose towards margin, viscid, red brown to bluish ochraceous brown, becoming paler ochraceous brown with age; often bicoloured, outer part hygrophanous and darker (grey) brown. Lamellae: crowded, edge even to crenulated; 4–7 mm broad, initially pale grey with a bluish tinge, later pale brown. Stipe: 4.5–8 (–11) × 1–1.7 cm, with a more or less distinct marginate bulb (→3.5) cm), ibrillose (not glossy like C. multiformis), whitish, but usually with a bluish tint at apex when (very) young, often becoming pale brown with age. Universal veil: white, sparse, sometimes viscid (at bulb margin). Context: in the pileus whitish, brownish below the cuticle, in the stipe apex with a bluish tint when (very) young. Smell: distinct of honey when cut, especially in the bulb. Spores: 8–9(–9.5) × 5–6 µm, MV = 8.6 × 5.3, Q = 1.64, ellipsoid(-ovoid) to more (sub)amygdaloid, sometimes broadly ellipsoid, medium strongly (to strongly) verrucose, warts small/dense to rather coarse and rounded, often somewhat conluent. Pileipellis: duplex, cutis thin, at surface of 2–3 µm wide, gelatinous, erect-entangled, hyaline hyphae. Subcutis/hypoderm tightly cemented, subcellular, with yellow brown-golden yellow parietal pigment, forming an amber-like embedment, some small brown, oleiferous spots present, and a few, more narrow zebra-striped, encrusted hyphae. DNA/ITS sequence: Eight of the 9 analysed sequences (including 5 generated here) are identical, only 1 differs by 1 nucleotide from the others. 193 Photo: Tor Erik Brandrud Fig. 17 – Cortinarius caesiolamellatus (Germany, TEB 428-10; +DNA) Scatter diagram: Geert Schmidt-Stohn Fig. 18 – Variation in spore size of C. caesiolamellatus and C. caesiophylloides. Each symbol = MV of spores measured per basidiome. 194 Ecology: In boreal-boreonemoral, montane and subalpine coniferous forests, often on rich, sometimes calcareous ground. In Europe apparently mainly/only under Picea, but in U.S.A. also found under Pinus. Distribution: Known mainly from C Europe and from S Fennoscandia, with a few outposts in SW Europe and N Fennoscandia. Widespread, but more precise distribution is uncertain due to confusion with C. caesiophylloides and C. multiformis. Also recorded from Washington, U.S.A. Collections examined: GERMANY. Baden-Württemberg, Schwaben, Ehingen a.d. Donau, Kohlhau, 28. Sept. 2010, G. Schmidt-Stohn, T. E. Brandrud, TEB 428-10 (herb. O) (DNA*); Kolhau, 28. Sept. 2010, G. Turrini (herb. Turrini) (DNA); Bockinhofer Holz, leg. K. Soop, KS-CO1938, (herb S) (DNA). Bavaria: Bad Hindelang, Oberjoch, 4. Oct. 2003, S. Garnica SG 0828 (herb. TUB) (DNA). Saarland: Schmelz, Hüttersdorf, 31. Oct. 2004, K. Montag (herb. TUB) (DNA). Thürringen: Thölendorf, 11.10.98, U. Luhmann UL 98/122 (herb. TUB 011841), GenBank No. AY669531 (as “C. allutus”) (DNA*). FRANCE. Herault: Bedarieux, TEB 22383, 242-83 (herb. O). ITALY. Trento, Santa Colomba, TEB 315-82 (herb. O). NORWAY. Oslo: Grorud, Steinbruvannet, 4 Sept. 2011, TEB 687-11 (herb. O) (DNA*). Vestfold, Tønsberg, TEB 278-80 (DNA*). Sogn & Fjordane: Lærdal, TEB 213-12 (herb. O) (DNA*). Oppland: Gran, Buhammeren, TEB 116-79 (herb. O). Lunner, Rustad, TEB 478-80 (herb. O). Lunner, S. Oppdalen , E. Bendiksen 10. Sept. 1978, TEB 72-88 (herb. O). Buskerud, Notodden, TEB 12911 (herb. O). Hedmark: Ringsaker, Brumunddal, TEB 171-80 (herb. O). FINLAND. Uusimaa, Espoo, Kunnarla, I. Kytövuori, IK94-852 (herb. H) (DNA). Perä-Pohjanmaa, Tervola, Peura, XXI Nordic Myc. Congr., 5. Sept. 2013, H. Lindström, E. Bendiksen, det. I. Kytövuori (herb. O; H). U.S.A. Washington, Olympic peninsula, Ocean Shore Park, 11. Nov. 2009, J. Ammirati & T. Niskanen, TN09-201 (herb. H), GenBank No. KF732571 (DNA*). Comments: Cortinarius caesiolamellatus and C. caesiophylloides are easily distinguished from related taxa in Multiformes by their initially bluish tinged lamellae and stipe apex. Based on material seen so far and available descriptions, these two blue-gilled taxa appear to be dificult to distinguish macromorphologically, but C. caesiolamellatus sees to have warmer, more redbrownish pileus colours, a more pronounced bicoloured pileus with a dark, hygrophanous outer part, and furthermore they differ in spore characters: C. caesiolamellatus has more ellipsoid, wider and more strongly verrucose spores. Finally, there seems to be a geographical differentiation in Europe; C. caesiolamellatus apparently being mainly a C. European taxon including southern parts of N Europe, whereas C. caesiophylloides is hitherto found only in Northern Europe. However, these taxa seem to be overlapping in morphological variation, and probably cannot always be identiied on morphological features. When all evidence is taken together (phylogenetically distinct, with <99% similarity + some morphological differentiation), it seems nevertheless reasonable to treat these as two species. More studies may reveal that these are possible to distinguish and recognize by macro/micromorphological characters. If the bluish tints have faded, C. caesiolamellatus can also be mixed with the conifer associated C. rufoallutus, C. multiformis and C. talimultiformis. The latter three, however, have less strongly ornamented spores and possess yellow brown, oleiferous pigment lumps and (C. rufoallutus) zebra-striped encrusted pigment in the hypodermium. Cortinarius caesiolamellatus is phylogenetically close to C. caesiophylloides, C. pallidorimosus and C. armenicorius, forming one of the two major subclades of sect. Multiformes. The species was described in Bidaud et al. (2006) both as C. rufoallutus var. caesiolamellatus and C. multiformis var. caesiophyllus, and their types conform 100% phylogenetically with our species (Figure 1, cfr. LiimaTainEn et al. 2014). Bluish variants probably referable to this taxon are otherwise mentioned by SchäFFEr (1949, sub nom. C. melliolens var. violascens ad. int.) and by kriEGlSTEinEr & GmindEr (2010, sub nom. C. allutus p.p.). Cortinarius multiformis variants with bluish lamellae and a distinct honey smell were distinguished 195 by the senior author as a separate taxon already in the late 1970ies, with the working name C. cyanoallutus (= C. caesiolamellatus). However, due to limited material and the complex variation in the C. multiformis complex, with only little micromorphological differences, this was never validly published (but introduced in Soop 2011 as C. multiformis var. cyanoallutus ined.). Bluish carpophores were not mentioned by MoSEr (1961), but he found that mycelium cultures of C. allutus could develop bluish colours. When later discussing the matter with M. Moser, he was of the opinion that presence/ absence of bluish pigments were not taxonomic relevant in the C. multiformis-C. allutus group. Then much later, bluish variants was irst formally published as C. rufoallutus var. caesiolamellatus (= C. multiformis var. caesiophyllus) by Bidaud et al. (2006), both covering the present C. caesiolamellatus. Cortinarius caesiophylloides Kytöv., Liimat., Niskanen, Brandrud & Frøslev Figs. 12, 18-20 Persoonia 33 (2014). Type: Finland. T. Niskanen 05-016, H6029792 (H, holotype). Icone: LiimaTainEn et al., Persoonia 33 (2014). Pileus: 4–8 cm broad, (hemi)spherical to plano-convex, viscid, centre with a dense pattern of small, darker, hygrophanous spots (pseudohygrophanous), towards margin with more elongated hygrophanous streaks/veins, or hygrophanous (water soaked) in outer zone; ochraceous brown to more clay brown, possibly also more fulvous brown, hygrophanous part somewhat darker umber to less vivid (grey) brown, becoming pale ochraceous brown-ochraceous yellow with age, even whitish ochre at centre. Lamellae: L = 60–80, crowded, edge even to crenulated; 4–7 mm broad,; initially pale grey with a bluish tinge, later pale brown. Stipe: 5–11 cm long, 1–1.5 cm, with a more or less distinct marginate bulb (→3.5 cm), ibrillose (not glossy like C. multiformis); whitish but usually, with a bluish tint at apex when (very) young, often becoming pale brown with age. Universal veil: white, sparse, sometimes viscid (at bulb margin). Context: white, brownish below the cuticle; slightly bluish at the apex of the stipe when (very) young. Smell: distinct of honey when cut, especially in the bulb. Spores: 8.5–9.5(‒10) × (4.5–)5–6 µm, (MV = 9.0 × 5.3 µm) Q = 1.70, (sub)amygdaloid, sometimes acutely amygdaloid, pale, weakly (to moderately) verrucose, warts dense, and often rounded-conluent. Pileipellis: duplex, epicutis with a gelatinous layer with sparse, erect-entangled, 2–3 µm wide, smooth, hyaline hyphae. Hypoderm present, of fairly thin-walled elements, with pale yellowish brown amber-like parietal pigment, some hyphae with yellow brown zebra-striped encrustations present, large yellow brown pigment lumps absent. DNA (ITS) sequence: All 6 ITS sequences available (including 3 analysed in the present study) are identical. Habitat: Boreal coniferous forests, associated mainly or only with Picea abies (one Finnish collection from dry Pinus forest apparently without Picea; LiimaTainEn et al. 2014). Mainly in richer low-herb types, sometimes on calcareous ground, once found in a rich sandy pine-spruce forest (TEB 720-13/DB-NO-181). Distribution: So far known only from Fennoscandia, with a core area probably in some northeastern regions (such as Trøndelag, Jämtland and N/C Finland); rare (known from seven localities). Collections examined: NORWAY. Nord-Trøndelag: Stjørdal, Beistadvollen, 13 Aug 2009, TEB 277-09 (herb. O) (DNA*). Buskerud: Ringerike, Eggemoen northeast, TEB 720-13, DBNO-181 (herb. O) (DNA*). Oppland: Lunner, TF 2006-112 (herb. C), GenBank No. KF732575 (DNA*). SWEDEN. Jämtland, Bräcke, Bodsjö, Sidsjö, I. Kytövuori 97-697 (herb. H) (DNA). 196 FINLAND. Kainuu: Paltamo, Oikarilankylä, Kivesvaara, I. Kytövuori, IK08-1554, H6032621 (herb. H), GenBank No. KF732574 (DNA*). Photo: Tor Erik Brandrud Fig. 19 – Cortinarius caesiophylloides Norway, TEB 277-09; +DNA) Photo: Tor Erik Brandrud Fig. 20 – Cortinarius caesiophylloides Norway, TEB 277-09; +DNA) 197 Comments: Cortinarius caesiophylloides is characterized by bluish tinges on the lamellae and stipe apex when (very) young, and a distinct honey smell in (bulb) context. However, these features the species share with C. caesiolamellatus, and these taxa may not always be possible to distinguish morphologically. The spore shape and ornamentation stand out as the best differential character(s): C. caesiophylloides has more strictly amygdaloid, somewhat narrower spores with usually dense and pale ornaments, whereas the spores of C. caesiolamellatus usually are more (broadly) ellipsoid, and with stronger and coarser/more conluent warts. Furthermore, the latter has a more vivid/warmer red brownish pileus colour, and a dark, hygrophanous outer zone. Phylogenetically, these are well-supported units, distinguished by 25 nucleotide differences in ITS regions. Cortinarius caesiophylloides is in fact phylogenetically closer to C. pallidirimosus (17 nucleotide differences) than to C. caesiolamellatus (Fig. 1, 2). These two taxa are also differentiated geographically; C. caesiophylloides appears to be a more northern species, hitherto only found in the boreal zone of Fennoscandia, and mainly in the northeastern taiga regions, whereas C. caesiolamellatus is more southern, with most inds from S Fennoscandia and montane C and S Europe. Their habitat preferences seem alike; they both occur mainly in richer Picea forests. With (i) distinct phylogenetical differentiation (<99% similarity), (ii) a geographical differentiation, and (iii) some morphological differentiation especially the spore shape, these qualify to the rank of species. Few collections have been studied fresh of these, and more material could throw light on some more possible macromorphological differences. The presence of two bluish European Multiformes taxa did not become apparent before extensive ITS sequencing of the group was performed, and this was published as C. caesiophylloides this year by LiimaTainEn et al. (2014). Without the aid of molecular methods, this rare C. caesiolamellatus look-a-like would probably not have been discovered. The bluish tinges on C. caesiolamellatus and C. caesiophylloides soon fade with exposure and age. When bluish tints are absent they can be mixed with C. multiformis and C. talimultiformis (see comments under C. caesiolamellatus). Ackowledgements Mika Bendiksby and Ingeborg Bjorvand Engh, University of Oslo, Gábor M. Kovács and his team at the Eötvös Loránd University, Budapest, Martin Unterseher, University of Greifswald and Ali Tahir and Marco Tines (Frankfurt am M.) are thanked for performing analyses on ITS sequences. The Cortinarius, Flora Photographica team is thanked for cooperation in the study of the species, as well as our Norwegian colleagues Egil Bendiksen (NINA), Katriina Bendiksen and Klaus Høiland (both University of Oslo), our Finnish colleagues Ilkka Kytövuori, Kare Liimatainen and Tuula Niskanen (University of Helsinki), and, inally our Italian collegaues Gianni Turrini and Claudio Rossi, who also have contributed with important photos. Klaus Høiland and Sigisfredo Garnica (University of Tübingen) are thanked for providing insight in yet unpublished molecular/ phylogenetic data. Furthermore, we thank all the mycologists that have provided material for molecular or morphological study (see Table 1 and collections examined). Finally, we thank the J.E.C. and its leaders Karl Kob and Oswald Rohner who has supported the work of our JEC-DNA group in many ways, including inancial support to sequence analyses. Literature Bidaud, a., moËnnE-loccoz, p., rEumaux, p. & x. carTErET (2009). 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