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The largest type study of Agaricales species to date: bringing identification and nomenclature of Phlegmacium (Cortinarius) into the DNA era

The largest type study of Agaricales species to date: bringing identification and nomenclature of Phlegmacium (Cortinarius) into the DNA era

Profile image of Balint DimaBalint DimaProfile image of Tobias FrøslevTobias FrøslevProfile image of Tuula NiskanenTuula Niskanen

Abstract Cortinarius is a species-rich and morphologically challenging genus with a cosmopolitan distribution. Many names have not been used consistently and in some instances the same species has been described two or more times under separate names. This study focuses on subg. Phlegmacium as traditionally defined and includes species from boreal and temperate areas of the northern hemisphere. Our goals for this project were to: i) study type material to determine which species already have been described; ii) stabilize the use of Friesian and other older names by choosing a neo- or epitype; iii) describe new species that were discovered during the process of studying specimens; and iv) establish an accurate ITS barcoding database for Phlegmacium species. A total of 236 types representing 154 species were studied. Of these 114 species are described only once whereas 40 species had one ore more synonyms. Of the names studied only 61 were currently represented in GenBank. Neotypes are proposed for 21 species, and epitypes are designated for three species. In addition, 20 new species are described and six new combinations made. As a consequence ITS barcodes for 175 Cortinarius species are released.

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Persoonia 33, 2014: 98 –140 www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE http://dx.doi.org/10.3767/003158514X684681 The largest type study of Agaricales species to date: bringing identification and nomenclature of Phlegmacium (Cortinarius) into the DNA era K. Liimatainen1, T. Niskanen 1, B. Dima!, I. Kytövuori 2, J.F. Ammirati 3, T.G. Frøslev 4 Key words Basidiomycota diversity DNA barcoding ITS taxonomy typification Abstract Cortinarius is a species-rich and morphologically challenging genus with a cosmopolitan distribution. Many names have not been used consistently and in some instances the same species has been described two or more times under separate names. This study focuses on subg. Phlegmacium as traditionally defined and includes species from boreal and temperate areas of the northern hemisphere. Our goals for this project were to: i) study type material to determine which species already have been described; ii) stabilize the use of Friesian and other older names by choosing a neo- or epitype; iii) describe new species that were discovered during the process of studying specimens; and iv) establish an accurate ITS barcoding database for Phlegmacium species. A total of 236 types representing 154 species were studied. Of these 114 species are described only once whereas 40 species had one ore more synonyms. Of the names studied only 61 were currently represented in GenBank. Neotypes are proposed for 21 species, and epitypes are designated for three species. In addition, 20 new species are described and six new combinations made. As a consequence ITS barcodes for 175 Cortinarius species are released. Article info Received: 2 July 2013; Accepted: 24 February 2014; Published: 8 September 2014. INTRODUCTION Studies using DNA sequence data have shown that the diversity of fungi far exceeds earlier expectations with many common species of macrofungi still to be described and named (Schmit & Mueller 2007). In addition, it is not always easy to determine which species have been described and which are new to science, unless the type specimens of existing names can be carefully studied, including DNA sequencing. For many species of macrofungi the names have been difficult or even impossible to interpret. The most difficult ones are the older names with brief descriptions and usually without type material. But even if type material exists it can be very difficult to be certain of the identification based only on morphology, especially in challenging genera like Cortinarius where there is considerable convergence in morphology, colouration, and microscopic features. Furthermore, the literature is often difficult to obtain, making it hard to get information on available names and their application by earlier taxonomists. Consequently, many names have not been used consistently and in some cases the same species has been described two or more times under separate names. In instances where there is no type material available, a neotype (or a lectotype if collections of the author are available) is required to stabilize the use of the name. Finally, old type collections that are considered historical materials may not be available for study or DNA sequencing requiring the selection of an epitype. 1 2 3 4 Department of Biosciences, Plant Biology, P.O. Box 65, FI-00014 University of Helsinki, Finland; corresponding author e-mail: kare.liimatainen@helsinki.fi. Botanical Museum, P.O. Box 7, FI-00014 University of Helsinki, Finland. Department of Biology, Box 351800, University of Washington, Seattle, WA 98195-1800, USA. Natural History Museum of Denmark, Center for Geogenetics, University of Copenhagen, Øster Voldgade 5–7, 1350 Københaven K, Denmark. The development of molecular techniques has provided a more unambiguous tool to identify species. Currently the most commonly used locus in species level taxonomy is the nuclear ribosomal internal transcribed spacer (ITS), which has been proposed as a universal barcode marker for fungi (Schoch et al. 2012). The region is present in several chromosomes and is arranged in tandem repeats that are thousands of copies long (Burnett 2003). Due to the high copy number the region usually is easy to amplify and sequence, even from very old specimens (Larsson & Jacobsson 2004). In Cortinarius the ITS was proposed as a species-identifier sequence already in 2007 by Frøslev et al. and in 2008 by Ortega et al. It has also been shown that in the majority of the cases ITS is suitable for species delimitation in Cortinarius. The results of the multi-gene phylogenetic study based on ITS, rpb1, and rpb2 regions by Frøslev et al. (2005) showed that inference from ITS alone is indicative of the species level phylogenetic delimitations of multi-gene analyses. Furthermore, the delimitations inferred from ITS usually correlate with the morphospecies (Frøslev et al. 2007, Ortega et al. 2008, Niskanen et al. 2012b). Cortinarius is the largest genus of Agaricales with a cosmopolitan distribution and over 2 000 described species (Kirk et al. 2008). Cortinarius species are important ectomycorrhizal fungi associated with different trees and shrubs, belonging to the order Fagales, families Caesalpiniaceae, Cistaceae, Diptero­ carpaceae, Myrtaceae, Pinaceae, Rhamnaceae, Rosaceae and Salicaceae as well as a few herbaceous plants in the Cypera­ ceae. Owing to their often narrow ecological preferences and sensitivity to environmental change, many Cortinarius species have been used as indicator species for valuable natural environments, e.g. in Sweden and Denmark (Vesterholt 1991, Hallingbäck & Aronsson 1998, Frøslev & Jeppesen 2011). Lately it also was suggested that they have a key role in the carbon cycling of boreal forests (Bödeker et al. 2011). © 2014 Naturalis Biodiversity Center & Centraalbureau voor Schimmelcultures You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium Many of the major studies in Cortinarius have dealt with North American and especially European species, while the species of southern hemisphere are somewhat less studied (Moser & Horak 1975, Garnica et al. 2002, Cleland 1976 (1935), Gasparini & Soop 2008). In Europe the most extensive studies have been done by Fries (e.g., 1821, 1836 –1838, 1851) from Sweden, Henry (e.g., 1951, 1958, 1981, 1986) and Bidaud et al. (e.g., 1992, 2010) from France, Moser (e.g. 1960, 1969 –1970, 1983) from Central Europe, Høiland (1984), Brandrud (e.g., 1996, 1998), Brandrud et al. (e.g., 1990, 2012) and Niskanen et al. (e.g., 2009, 2011a, 2013a) mainly from northern Europe, Frøslev et al. (e.g., 2006, 2007) from northern and Central Europe, Orton (1955, 1958) from Great Britain, and Ortega et al. (2008) and Suárez-Santiago et al. (2009) from mediterranean area. Selected papers of some of the major contributors to Cortinarius systematics in North America include Peck (1873; also see Gilbertson 1962), Kauffman (1918, 1923, 1932), Smith (1939, 1942, 1944), Ammirati (1972), Ammirati et al. (2013), Garnica et al. (2009), Liu et al. (1997), Moser & Ammirati (1996, 1999), Moser et al. (1995), Bojantchev (2011a, b) and Niskanen et al. (2013b, c). Very little is known about the distribution of Cortinarius species on a larger scale or the differences in the species composition between the continents, although species in the southern hemisphere are distinct from those in the northern hemisphere (Peintner et al. 2004, Garnica et al. 2005). However, recent molecular studies on Cortinarius have shed some light on these questions for North America and Europe (Moser & Peintner 2002, Matheny & Ammirati 2006, Garnica et al. 2009, 2011, Harrower et al. 2011, Ammirati et al. 2013, Niskanen et al. 2011b, 2012c, 2013b, c). These studies show several patterns of species distributions. There are species common to North America and Europe, especially those species from more northern and montane conifer forests, i.e. Cortinarius aureofulvus M.M. Moser s.auct., C. napus Fr. and C. pinophilus Soop, but also endemic species occur both in western North America, eastern North America and Europe, i.e. C. elegantio­occidentalis Garnica & Ammirati in western North America, C. hesleri Ammirati, Niskanen, Liimat. & Matheny in eastern North America and C. puniceus P.D. Orton in Europe. Cortinarius species composition is somewhat similar between eastern North America and Europe, but there appears to be less similarity between Europe and western North America. Different infrageneric classification systems have been proposed for Cortinarius, i.e. Brandrud et al. (1990) divided the genus in four subgenera: Cortinarius, Myxacium, Phlegmacium and Telamonia. Recent molecular analyses have shown that Cortinarius is monophyletic, but also that many of the infrageneric groups such as subg. Phlegmacium are not monophyletic (Peintner et al. 2004, Garnica et al. 2005). Nonetheless, several authors have treated subg. Phlegmacium in similar ways (Moser 1983, Bidaud et al. 1994, Brandrud et al. 2012) including in it species with viscid to glutinous pileus and dry, often bulbous stipe, or dry-capped, stout species with a yellow KOH reaction, i.e. the species of sect. Phlegmacioides (Niskanen et al. 2012a). Molecular techniques have been in use for more than a decade, and the sequencing of ITS regions even from older Cortinarius specimens is possible. Furthermore, type studies are essential for a stable and consistent application of names in Cortinarius where currently a large percentage of the Cortinarius sequences are incorrectly named or without a name in the public sequence databases (e.g., Niskanen et al. 2011a). While several papers present sequences for individuals or groups of species, for example Garnica et al. (2009) and Niskanen et al. (2011a), the only large study is that of Frøslev et al. (2007) where 52 types of Cortinarius sect. Calochroi were sequenced. The present 99 study focuses on Cortinarius subg. Phlegmacium as traditionally defined and includes species from boreal and temperate areas of the northern hemisphere. Our goals for this project are to: i) study type material to determine which taxa have already been described; ii) stabilize the use of Friesian and other older names by choosing a neo- or epitype; iii) describe new species that were discovered during the process of studying specimens; and iv) establish an accurate ITS barcoding database for Phlegmacium species. MATERIALS AND METHODS Taxon sampling The type specimens of species of subg. Phlegmacium published over many years by J.F. Ammirati, A. Bidaud, T.E. Brandrud, G. Chevassut, J. Favre, R. Henry, P.A. Karsten, C.H. Kauffman, R. Kühner, K.H. McKnight, M.M. Moser, P. Moënne-Loccoz, P. Reumaux, A.H. Smith and H.D. Thiers were sampled as well as relevant collections published and illustrated in Brandrud et al. (1990, 1992, 1994, 1998). The species of sections Calochroi, Fulvi and Riederi described from Europe were generally excluded. The first two have been treated by Frøslev et al. (e.g., 2007) and the latter will be treated by Brandrud et al. in the near future. In some instances type material could not be acquired from herbaria, for example, the Cortinarius type collections of C.H. Peck were only recently available on loan from the New York State Museum and will be included in a later study. Our aim was to have at least two sequences per species in our study. Therefore, in addition to sequences from type specimens, either our own unpublished sequences or additional sequences retrieved from the sequence databases GenBank and UNITE were included. Unpublished sequences were also supplied by D. Bojantchev, K. Hughes and A. Taylor. Information on the sequences of type specimens is available in Table 1 and information on other sequences included in the phylogenetic analysis is available in Table 2. Herbarium acronyms follow Index Herbariorum (Thiers 2013). Molecular analyses DNA was extracted from a few milligrams of dried material (a piece of lamella) with the NucleoSpin Plant kit (MachereyNagel, Düren, Germany), or with various CTAB protocols in Brandrud’s and Frøslev’s specimens (see Frøslev et al. 2005, 2007). Primers ITS 1F and ITS 4 (White et al. 1990, Gardes & Bruns 1993) were used to amplify ITS regions. The same primer pairs were used in direct sequencing. For problematic material the primer combinations ITS 1F/ITS 2 and ITS 3/ITS 4 were also used. PCR amplifications were performed in a 25 µL reaction mix with about 70 ng extracted DNA, 1 U Phusion High-Fidelity DNA polymerase and 1× HF buffer (Finnzymes), 200 mM of each dNTP and 0.5 μM of each primer. The PCR reactions were run on a MBS 0.2 G Thermal Cycler (Thermo Hybaid) with the following settings: denaturation for 30 s at 98 °C, followed by 35 cycles of denaturation for 10 s at 98 °C, annealing for 30 s at 50 °C and extension for 30 s at 72 °C. The PCR products were purified using an ExoSAP-IT purification kit (Amersham Biosciences). Sequencing was performed on both strands using a BigDye Terminator v. 1.1 Sequencing kit (Applied Biosystems). Reactions were performed in 10 µL with 1 µL of PCR product, 1.3 mM of primer (ITS 1F or ITS 4), 1 µL 5X sequencing buffer, and 1 µL of Terminator Ready Reaction Mix. Reactions were run for 1 min at 96 °C, followed by 30 cycles of 30 s at 96 °C, 15 s at 50 °C and 4 min at 60 °C. Unincorporated dye terminators and primers were removed by Sephadex G-50 DNA Grade Fine (Amersham Biosciences) purification system, (text continues on p. 117) Species Voucher Herb. Locality Ecology Current name Collection date Collector 100 Table 1 Type specimens studied. Sequences of the type materials of C. caesiocinctus, C. cobaltinus, C. gracilior, C. mahiquesii, C. norrlandicus and C. van­campiae retrieved from the GenBank or UNITE. Short ITS sequences excluded from the analysis marked with *. GenBank number C. acidophilus Brandrud 1997 (holotype) TEB61-79 O Norway, Oppl, Lunner, Østäsen In forest of Picea abies C. pseudonaevosus Rob. Henry 1957 01.08.1979 T.E. Brandrud KF732241 C. acystidiosus Thiers 1960 (holotype)* 1902 MICH USA, Texas, San Jacinto Co., Sam Houston National Forest, Coldspring In pine-hardwood forest C. acystidiosus Thiers 1960 23.05.1953 H.D. Thiers KF732242 C. aggregatus Kauffman 1918* MICH10311 MICH USA, Michigan, Jackson, Vandercook Park, Jackson In low woods C. aggregatus Kauffman 1918 09.09.1907 C.H. Kauffman KF732243 C. albescens A.H. Sm. 1944 (holotype) 17522 MICH USA, Washington, Olympic National Park, Olympic Hot Springs Under conifers C. albescens A.H. Sm. 1944 02.10.1941 A.H. Smith KF732244 C. albofragrans Ammirati & M.M. Moser 1997 (holotype) 95/595 IB USA, California, Del Norte Co., Highway 199, Danger Point Under Quercus, also evergreen oaks (Q. vacci­ C. albofragrans Ammirati & M.M. Moser niifolia, Q. chrysolepis, Lithocarpus densi­ 1997 flora) with some Pseudotsuga menziesii 20.11.1996 M. Moser KF732245 C. alnobetulae Kühner 1989 (syntype) 53-6 G France Under Alnus viridis C. alnobetulae Kühner 1989 Unknown Unknown KF732246 C. alticaudus Reumaux 2008 (holotype) PML1632 PC France, Lozère, bois de la Sagne In Pinus forest C. alticaudus Reumaux 2008 30.10.1989 P. Reumaux KF732247 C. amarocaerulescens Bidaud 2009 (holotype) AB99-11-362 PC France, Loire, Montbrison In mixed forest C. infractus (Pers.: Fr.) Fr. 1838 09.11.1999 A. Bidaud KF732248 C. amnicola A.H. Sm. 1942 (holotype) 15381 MICH USA, Michigan, Ann Arbor Under butternut, walnut, and bassword on low ground along streams C. amnicola A.H. Sm. 1942 15.09.1940 A.H. Smith KF732249 C. anfractoides Rob. Henry & Trescol 1987 (holotype)* RH1898 PC France In frondose forest under Quercus ilex C. anfractoides Rob. Henry & Trescol 1987 11.1984 G. Chevassut & F. Trescol KF732250 C. anfractoides var. cinereoclarus Bidaud 2009 (holotype) AB08-09-155 PC France, Ain, Cerin In calcareous deciduous forest C. persoonianus Bidaud 2009 07.09.2008 A. Bidaud KF732251 KF732252 C. arenicola A.H. Sm. 1942 (holotype) 15315 MICH USA, Michigan, Waterloo, Waterloo Project Under Sassafras in dry sandy open woods C. arenicola A.H. Sm. 1942 12.09.1940 A.H. Smith C. areni­silvae (Brandrud) Brandrud 2012 (holotype) CFP461b S Sweden, Ång, Graninge, Viksmon In boreal coniferous forest with Pinus and Picea on sandy soil C. areni­silvae (Brandrud) Brandrud 2012 25.08.1986 T.E. Brandrud et al. KF732253 C. argutipes Bidaud & Reumaux 1996 (holotype)* 713 G France, Creuse Under Fagus on acid soil C. chromataphilus Rob. Henry 1989 15.10.1987 D. Brion KF732254 C. atrochalybaeus M.M. Moser & Ammirati 2000 (holotype) 95/630 IB USA, California, Del Norte Co., Highway 199, Smith River Middle Fork, Danger Point Under Lithocarpus densiflora, Arbutus menziesii, Quercus vacciniifolius and Q. chrysolepis on calcareous soil C. atrochalybaeus M.M. Moser & Ammirati 2000 29.11.1995 J. Ammirati KF732255 C. aurantionapus Bidaud & Reumaux 2006 (holotype)* PML4893 PC France, Drôme, Romeyer Under Picea and Pinus sylvestris on calcareous soil C. talus Fr. 1838 09.10.1992 J. Garin KF732256 C. aurantionapus var. similis Moënne-Locc. 2006 (holotype) PML883 PC France, Haute-Savoie, Avernioz Under Picea on calcareous soil C. talimultiformis Kytöv., Liimat., Niskanen, A.F.S. Taylor & Sesli sp. nov. 06.06.1988 P. Moënne-Loccoz KF732257 AB05-11-404 PC France, Ardèche, Lagorce Under Quercus ilex on calcareous soil C. aurantiopallidus Bidaud 2006 11.11.2005 A. Bidaud KF732258 53/10 IB Germany, Schwenningen In coniferous forest C. badiolatus (M.M. Moser) M.M. Moser 1967 28.08.1953 H. Haas KF732259 C. balteatialutaceus Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK09-751 H Sweden, Jmt, Frostviken, Jormlien, Säterklumpen In Betula forest with solitary Picea C. balteatialutaceus Kytöv., Liimat. & Niskanen sp. nov. 09.07.2009 P. & I. Kytövuori KF732586 C. balteatibulbosus Kytöv., Niskanen, Liimat., Bojantchev & A.F.S. Taylor sp. nov. (holotype) IK98-1624 (H6033539) H Finland, U, Espoo, Nuuksio, Pirttimäki In half-open cut meadow C. balteatibulbosus Kytöv., Niskanen, 04.09.1998 Liimat., Bojantchev & A.F.S. Taylor sp. nov. I. Kytövuori KF732589 C. balteaticlavatus Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK96-595 (H6032412) H Finland, PH, Virrat, Hauhuu, Sikosaari, Salmela On the grassy roadside with young Picea, Betula, Populus tremula, Alnus incana and Salix spp. C. balteaticlavatus Kytöv., Liimat. & Niskanen sp. nov. 23.08.1996 I. Kytövuori KF732596 C. balteatoalbus Rob. Henry 1985 (isotype)* RH82.98 PC France In submontane Picea forest C. balteatoalbus Rob. Henry 1985 Unknown Unknown KF732260 C. balteatotomentosus Rob. Henry ex Rob. Henry 1985 (holotype) RH306 PC France, Doubs, Boujaille In montane forest (mostly) of Picea abies C. balteatus (Fr.) Fr. 1838 Unknown Unknown KF732261 C. balteatus (Fr.) Fr. 1838 (neotype) CFP940 S Sweden, Ång, Säbrå, Överdal Under Picea in cultivated area C. balteatus (Fr.) Fr. 1838 03.08.1990 T.E. Brandrud et al. KF732262 C. balteatus var. praestantoides Reumaux 1996 (holotype) 760 G France, Ile-de-France, Forêt de Rambouillet, Etang d’Or In shrubs in Quercus forest C. flavescentipes Reumaux 1996 23.10.1982 P. Reumaux KF732263 C. barrentium Poirier & Reumaux 1993 (holotype)* 2398 G France, Loiret, Arboretum des Barres Under conifers C. barrentium Poirier & Reumaux 1993 13.11.1991 J. Poirier KF732264 Persoonia – Volume 33, 2014 C. aurantiopallidus Bidaud 2006 (holotype) C. badiolatus (M.M. Moser) M.M. Moser 1967 (holotype)* 45385 MICH USA, Idaho, Seven Devils Mts, Heaven’s Gate Ridge Under conifers C. bigelowii Thiers & A.H. Sm. 1969 26.07.1954 H.E. Bigelow KF732265 C. boreicyanites Kytöv., Liimat., Niskanen & A.F.S. Taylor sp. nov. (holotype) CFP931 S Sweden, Jmt, Ragunda, Böle In birch forest on rich ground (Betula, Picea) C. boreicyanites Kytöv., Liimat., Niskanen & A.F.S. Taylor sp. nov. 07.24.1990 T.E. Brandrud et al. KF732296 C. boreidionysae Kytöv., Liimat., Niskanen & Dima sp. nov. (holotype) IK97-1220 H Finland, PeP, Tervola, Peura, Raemäki In grass-herb-spruce forest with springfed depressions, on calcareous ground C. boreidionysae Kytöv., Liimat., Niskanen & Dima sp. nov. 11.09.1997 I. Kytövuori KF732488 C. borgsjoeensis Brandrud 1992 (isotype) CFP728 S Sweden, Jmt, Ragunda, Kullstabodarna In herbaceous spruce forest C. borgsjoeensis Brandrud 1992 31.08.1988 T.E. Brandrud et al. KF732266 C. brunneiaurantius Kytöv., Liimat. & Niskanen sp. nov. (holotype) JV17979 (H6032422) H Finland, V, Turku, Ruissalo, Kansanpuisto Tilia alley, also Quercus robur nearby, on clayey-mull soil C. brunneiaurantius Kytöv., Liimat. & Niskanen sp. nov. 22.09.2001 J. Vauras KF732600 C. brunneolividus Bidaud 1996 (holotype) 3734 G France, Isère, Optevoz In calcareous deciduous forest of Quercus and Carpinus C. brunneolividus Bidaud 1996 14.09.1993 C. Blanc KF732268 C. brunneoviolaceus Bidaud 1996 (holotype) 2951 G France, Isère, Arzay Under Castanea on acid soil C. brunneolividus Bidaud 1996 01.10.1992 A. Bidaud KF732269 C. cacodes M.M. Moser & Ammirati 2000 (holotype) 91/618 IB USA, California, Mendocino, Russian Gulch State Park In mixed coniferous forest with Tsuga, Pseudotsuga, Abies C. cacodes M.M. Moser & Ammirati 2000 30.11.1991 M. Moser KF732270 C. caerulescens (Schaeff.) Fr. 1838 (epitype) CFP853 S Belgium, Brabant, Tervuren In beech forest on calcareous ground C. caerulescens (Schaeff.) Fr. 1838 23.09.1989 T.E. Brandrud et al. KF732271 C. caesiocinctus Kühner 1989 (holotype) 57-13 G France, Haute-Savoie, Le Môle In mixed forest with Fagus and Picea C. caesiocinctus Kühner 1989 30.08.1957 R. Kühner DQ663239 C. caesiocolor Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK00-029 H Finland, U, Lohja, Jalassaari, Tamminiemi, by a track With Betula, Populus tremula, Quercus, Corylus C. caesiocolor Kytöv., Liimat. and Salix caprea, on calcareous ground & Niskanen sp. nov. 27.08.2000 I. Kytövuori KF732603 C. caesiophylloides Kytöv., Liimat., Niskanen, Brandrud & Frøslev sp. nov. (holotype) TN05-016 (H6029792) H Finland, ES, Joutsa, Koivuranta In fairly young, mesic to damp, Picea abiesdominated forest with some Betula and Pinus sylvestris C. caesiophylloides Kytöv., Liimat., Niskanen, Brandrud & Frøslev sp. nov. 30.08.2005 K. Liimatainen & T. Niskanen KF732572 C. calojanthinus M.M. Moser & Ammirati 1999 (holotype) 97/220 IB USA, Wyoming, Teton National Forest, Calypso Creek, Flagstaff Road In subalpine forest under Picea engelmannii, Abies lasiocarpa C. calojanthinus M.M. Moser & Ammirati 1999 21.08.1997 M. Moser & J. Ammirati KF732272 C. calyptratus A.H. Sm. 1939 (holotype) 8352 MICH USA, California, Crescent City Under mixed spruce and redwood, presumably also Lithocarpus and Quercus C. calyptratus A.H. Sm. 1939 03.11.1937 A.H. Smith KF732273 C. calyptrodermus A.H. Sm. 1942 (holotype) 15356 MICH USA, Michigan, Sharron Hollow In low woods of second growth oak and basswood (Tilia) and various shrubs C. calyptrodermus A.H. Sm. 1942 14.09.1940 A.H. Smith KF732274 C. castaneicolor A.H. Sm. 1944 (holotype) 17926 MICH USA, Washington, Olympic National Park, Olympic Hot Springs Under conifers C. castaneicolor A.H. Sm. 1944 15.10.1941 A.H. Smith KF732275 C. cephalixoides M.M. Moser & Thiers 1995 (holotype) 87/188 IB USA, Wyoming, Teton National Forest, Flagstaff Road In subalpin forest under Picea engelmannii C. cephalixoides M.M. Moser & Thiers 1995 09.08.1987 H.D. Thiers KF732276 C. cephalixolargus Rob. Henry 1977 (holotype) 6048 PC France, Bois de Dampierre In mixed frondose forest under Fagus, Quercus, Carpinus and Betula C. largus Fr. 1838 Unknown R. Henry KF732277 C. chromataphilus Rob. Henry 1989 (holotype) 86.90 PC France, Luxeuil In mixed forest C. chromataphilus Rob. Henry 1989 1986 Exposition KF732278 C. cinctipes Bidaud, Eyssart. & Hermitte 2004 (holotype) GE 02-100 PC France, Var, Les Mayons Under Quercus suber and Erica arborea C. pseudocephalixus Bidaud & Moënne-Locc. 2000 01.11.2002 J.-C. Hermitte KF732279 C. citrinifolius A.H. Sm. 1939 (holotype) 3158 MICH USA, Washington, Olympic National Park, Boulder Creek Under fir C. citrinifolius A.H. Sm. 1939 15.10.1935 A.H. Smith KF732280 C. citrinipedes A.H. Sm. 1942 (holotype) 15305 MICH USA, Michigan, Ann Arbor On humus in oak woods C. citrinipedes A.H. Sm. 1942 11.09.1940 A.H. Smith KF732281 C. citriolens Ammirati & M.M. Moser 1999 (holotype) 97/122 IB USA, Wyoming, Teton National Forest Flagstaff Creek In subalpine forest under Picea engelmannii, Abies lasiocarpa C. citriolens Ammirati & M.M. Moser 1999 06.08.1997 M. Moser & J. Ammirati KF732282 C. claricolor (Fr.) Fr. 1838 (neotype) CFP691 S Sweden, Ång, Stigsjö, Uland, Langmyrberget In spruce forest with blueberry C. claricolor (Fr.) Fr. 1838 09.08.1988 T.E. Brandrud et al. KF732283 C. clarobaltoides var. longispermus Reumaux 1996 (holotype) 833 G France, Ile-de-France Under conifers C. clarobaltoides var. longispermus Reumaux 1996 15.09.1987 M. Pelerin K. Liimatainen et al.: Identification and nomenclature of Phlegmacium C. bigelowii Thiers & A.H. Sm. 1969 (holotype) KF732284 4038 G France, Yonne, Forêt de Hervaux In frondose forest C. largus Fr. 1838 04.10.1994 Mlle Maité KF732285 2932 G France, Ardennes, Forêt de Belval Under Quercus and Fagus on clayeycalcaerous soil C. largus Fr. 1838 19.09.1992 P. Reumaux & F. Reumaux KF732286 C. cobaltinus Kytöv., Liimat. & Niskanen 2013 (holotype) TN02-1012 (H6032404) H Finland, Ks, Kuusamo, Oulanka National Park In herb-rich Picea abies forest with some Pinus, Betula and Populus on calcareous ground C. cobaltinus Kytöv., Liimat. & Niskanen 2013 22.09.2002 T. Niskanen, I. Kytövuori & K. Liimatainen KF673470 C. collocandoides Reumaux 2009 (holotype) PML5087 PC France, Yvelines, étang d’Or, Rambouillet In mixed forest C. collocandoides Reumaux 2009 27.10.1996 G. Redeuilh & P. Reumaux KF732287 C. concrescens Secr. ex Bidaud, Moënne-Locc. & Reumaux 1996 (holotype) 3578 G France, Haute-Savoie, Le Danay, St-Jean-de-Sixt Under Picea abies and Alnus viridis C. balteatoalbus Rob. Henry 1985 10.10.1993 M. Mugnier KF732288 101 C. clarus Reumaux 1996 (holotype) C. claviceps Reumaux 1996 (holotype) 102 Table 1 (cont.). Species Voucher Herb. Locality Ecology Current name Collection date Collector GenBank number C. congeminus Moënne-Locc. & Reumaux 1995 (holotype) 3422 G France, Ardennes, Forêt de Belval In deciduous forest on clayey-calcareous soil C. largus Fr. 1838 03.10.1992 P. Reumaux & F. Reumaux KF732289 C. corrugis A.H. Sm. 1944 (holotype) 16842 MICH USA, Washington, Baker National Forest, Anderson Lookout, Ermine Creek Trail Under conifers C. turmalis Fr. 1838 11.09.1941 A.H. Smith KF732290 C. crassus Fr. 1838 (neotype) CFP938 S Sweden, Ång, Säbrä, Hårsta In wet mixed forest with Sphagnum and Pinus, Picea, Betula C. crassus Fr. 1838 28.07.1990 T.E. Brandrud et al. KF732291 C. cremeiamarescens Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK11-014 H Sweden, Gtl, Alskog and När parish Mesic to damp spruce forest with some Pinus, Quercus and Corylus C. cremeiamarescens Kytöv., Liimat. & Niskanen sp. nov. 27.09.2011 I. Kytövuori KF732493 C. crenulatus Rob. Henry ex Bidaud & Reumaux 2006 (holotype) PML4866 PC France, Allier, Forêt de Tronçais In deciduous forest C. talus Fr. 1838 17.10.1992 F. Lopez KF732292 C. cruentipellis Kytöv., Liimat., Niskanen & Dima sp. nov. (holotype) TN03-1451 H Sweden, Öl, Långlöt, Åstad, Nitares hägn Grassy pasture with Corylus and Juniperus C. cruentipellis Kytöv., Liimat., Niskanen & Dima sp. nov. 13.09.2003 I. Kytövuori, K. Liimatainen & T. Niskanen KF732539 KF732293 C. cumatilis Fr. 1838 (neotype) IK98-2164 H Sweden, Nrk, Hidinge, Garphyttan Spruce forest C. cumatilis Fr. 1838 20.09.1998 I. Kytövuori C. cupreorufus Brandrud 1994 (isotype) CFP1038 S Sweden, Upl, Älvkarleby, Billudden In dry spruce forest on rich ground C. cupreorufus Brandrud 1994 03.10.1990 T.E. Brandrud et al. KF732294 C. cupreoviolaceus Bidaud & Reumaux 1996 (holotype) 3426 G France, Ardennes, Forêt de Belval In deciduous forest on clayey-calcareous soil C. largus Fr. 1838 27.09.1992 P. Reumaux & F. Reumaux KF732295 AT2005069 H Sweden, Upl, Uppsala, Stadsskogen In mixed forest C. cyanites Fr. 1838 26.08.2005 A. Taylor KF732355 RH8673 PC France, Paris region In frondose forest C. delaportei Rob. Henry 1988 Unknown A. Delaporte KF732297 C. dionysae var. avellaneus Rob. Henry ex Bidaud & Carteret 2008 (holotype) AB97-10-361 PC France, Ain, Champdor In mixed calcareous forest C. dionysae var. avellaneus Rob. Henry ex Bidaud & Carteret 2008 21.10.1997 G. Chamonaz KF732298 C. eliae Bidaud, Moënne-Locc. & Reumaux 1996 (holotype) 1032 G France, Haute-Savoie, Vieugy In herbaceous Quercus forest C. eliae Bidaud, Moënne-Locc. & Reumaux 1996 27.10.1988 P. Moënne-Loccoz KF732299 C. elotoides M.M. Moser & McKnight 1995 (holotype) 87/60 IB USA, Wyoming, Teton National Forest, Turpin Meadow Under Picea pungens, P. engelmannii, Pseudotsuga menziesii in subalpine forest C. elotoides M.M. Moser & McKnight 1995 23.07.1987 Unknown KF732300 C. eumarginatus Rob. Henry ex Bidaud, Carteret & Reumaux 2009 (holotype) AB07-10-175 PC France, Ain, Chanay, col de Richemont In coniferous forest C. purpurascens Fr. 1838 16.10.2007 A. Bidaud KF732301 C. evosmus Joachim ex Bidaud & Reumaux 2006 (holotype)* PML4837 PC France, Oise, Coye-la-Forêt In deciduous forest on calcareous soil C. evosmus Joachim ex Bidaud & Reumaux 2006 20.10.1995 M. Diamond KF732302 C. flavaurora M.M. Moser & McKnight 1995 (holotype)* 89/187 IB USA, Wyoming, Teton National Forest, Fourmile meadow Under Picea engelmannii in subalpine forest C. elotoides M.M. Moser & McKnight 1995 07.08.1989 Unknown KF732303 C. flavescentipes Reumaux 1996 (holotype) 3606 G France, Loiret, Forêt de Montargis Under Quercus C. flavescentipes Reumaux 1996 27.10.1993 Participants of KF732304 mycological field trip in Sully-sur-Loire C. flavipallens Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK08-1729 (H6032745) H Finland, Kn, Kajaani, Hietalahti In fairly damp grass-herb-spruce forest with Pinus, Betula and Populus tremula, on calcareous ground C. flavipallens Kytöv., Liimat. & Niskanen 13.09.2008 sp. nov. I. Kytövuori KF732554 C. flavivelatus Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK98-885 H Sweden, Nb, Pajala, Junosuando In dryish Picea abies heath forest with Pinus, Betula, and open meadows C. flavivelatus Kytöv., Liimat. & Niskanen 15.08.1998 sp. nov. I. Kytövuori KF732528 C. flavobulbus Ammirati & M.M. Moser 1997 (holotype) 95/ 629 IB USA, California, Del Norte Co., Highway 199, Danger Point Under Quercus vacciniifolia and Q. garrayana in rather dry habitats on basic soils C. flavobulbus Ammirati & M.M. Moser 1997 29.11.1995 M. Moser KF732305 C. foetens (M.M. Moser) M.M. Moser 1967 (holotype) 51/128 IB Austria, Tyrol, Hötting-Innsbruck, Buchtal In mixed deciduous forest under Fagus C. foetens (M.M. Moser) M.M. Moser 1967 13.09.1951 M. Moser KF732306 C. fraudulosus Britzelm. 1885 (neotype) IK95-1852 S Germany, Baden-Württemberg, Freudenstadt, Heiligenbronn In gently sloping grass-herb coniferous forest (Abies alba, Picea abies, Fagus sylvatica) on calcareous ground C. fraudulosus Britzelm. 1885 05.10.1995 I. Kytövuori KF732518 C. fraudulosus var. patrickensis M.M. Moser 2000 (holotype) 95/ 617 IB USA, California, Humboldt Co., Trinidad, Patrick’s Point State Park Under Picea sitkensis and Pseudotsuga menziesii C. patrickensis (M.M. Moser) Niskanen, Liimat., Kytöv., Bojantchev & Ammirati comb. nov. 25.11.1995 M. Moser KF732307 C. frondosophilus Bidaud 2001 (holotype) PML4817 PC France, Ain, Cerin In deciduous forest on calcareous soil C. platypus (M.M. Moser) M.M. Moser 1967 16.11.1997 A. Bidaud KF732562 Persoonia – Volume 33, 2014 C. cyanites Fr. 1838 (neotype) C. delaportei Rob. Henry 1988 (holotype) 91/ 682 IB USA, California, Mendocino, Caspar Little Lake Road In coniferous forest with Pseudotsuga, Tsuga and Sequoia C. fuligineofolius (M.M. Moser) M.M. Moser & Peintner 2002 06.12.1991 M. Moser KF732308 C. fulmineus var. sulphureus Kauffman 1918 (lectotype)* MICH10351 MICH USA, Michigan, Washtenaw, Ann Arbor On the ground among humus, in frondose or mixed wood C. fulmineus var. sulphureus Kauffman 1918 03.10.1910 C.H. Kauffman KF732309 C. gentianeus Bidaud 1993 (holotype) 2575 G France, Ain, Lampnas In calcareous Fagus forest with Betula C. gentianeus Bidaud 1993 08.10.1991 M. Russi KF732310 C. genuinus Rob. Henry ex Bidaud & Carteret 2009 (holotype) XC2005-132 PC France, Seine-Maritime, Les Petites Dalles, Valmont In deciduous forest C. collocandoides Reumaux 2009 02.09.2005 X. Carteret KF732311 C. georgiolens Rob. Henry 1986 (holotype)* RH3153 PC France, Languedoc-Roussillon In forest of Quercus ilex C. georgiolens Rob. Henry 1986 Unknown Unknown KF732312 C. glaucocephalus M.M. Moser, Ammirati & Halling 2000 (holotype) 95/ 679 IB USA, California, Mendocino Co., Caspar Little Lake Rd. In mixed forests with Tsuga, Pseudotsuga, Pinus ponderosa, Abies and Arctostaphylos manzanita C. glaucocephalus M.M. Moser, Ammirati & Halling 2000 07.12.1995 J. Ammirati & M. Moser KF732313 C. glaucopoides Kauffman 1923 (holotype) MICH10358 KF732314 MICH USA, Colorado, Grand, Leal Under spruce and fir C. glaucopus (Schaeff.: Fr.) Gray 1821 16.08.1917 C.H. Kauffman C. glaucopus (Schaeff.: Fr.) Gray 1821 (neotype) CFP786 S Sweden, Mpd, Alnö, Ås brygga In dry spruce forest on calcareous ground C. glaucopus (Schaeff.: Fr.) Gray 1821 21.09.1988 T.E. Brandrud et al. KF732315 C. glaucopus var. olivaceus f. ingratus Moënne-Locc. 2008 (holotype) PML762 PC France, Haute-Savoie, forêt de la Semine, Clarafond In calcareous deciduous forest C. scaurocaninus Chevassut & Rob. Henry 1982 05.11.1987 P. Moënne-Loccoz KF732316 C. glaucopus var. subrubrovelatus Bidaud 2008 (holotype) AB97-11-431 PC France, Ain, Farges, in mixed forest with Populus tremula, Corylus sativa, Fagus sylvatica, Abies alba In deciduous forest C. subrubrovelatus (Bidaud) Kytöv., Liimat., Niskanen & Dima comb. nov. 01.11.1997 A. Bidaud KF732317 C. gracilior (Jul. Schäff. ex M.M. Moser) M.M. Moser 1967 (holotype) 58/ 73 IB Germany, Baden-Württemberg, Randengebiet near Zollhaus In Fagus forest on calcareous soil C. gracilior (Jul. Schäff. ex M.M. Moser) M.M. Moser 1967 09.10.1958 M. Moser UDB001082 C. gratus Reumaux 2008 (holotype) PML85 PC France, Haute-Savoie, forêt de la Mandallaz, Choisy In deciduous forest under Fagus C. leonicolor Reumaux 2001 27.09.1986 P. Moënne-Loccoz KF732318 C. griseocoeruleus Ammirati & M.M. Moser 1997 (holotype) 95/ 685 IB USA, California, Mendocino Co., about 8 km east of Mendocino on road 408 Under Lithocarpus densiflora Moser 1997 C. griseocoeruleus Ammirati & M.M. 08.12.1995 M. Moser KF732319 KF732320 C. herculeolens Bidaud 1996 (holotype)* 3700 G France, Montbrison, Loire Under Quercus petraea on basaltic soil C. chromataphilus Rob. Henry 1989 11.11.1992 A. Bidaud C. herpeticus Fr. 1838 (neotype) CFP936 S Sweden, Ång, Säbrå, Hällenyland Under Picea on cultivated area C. herpeticus Fr. 1838 20.07.1990 T.E. Brandrud et al. KF732321 C. hysginicolor Bidaud 1995 (holotype)* 2955 G France, Ain, Innimont In young herbaceus, calcareous Picea forest C. badiolatus (M.M. Moser) M.M. Moser 1967 03.10.1992 A. Bidaud KF732322 C. immixtus Kauffman 1932 (holotype)* MICH10365 MICH USA, Washington, Mason, Lake Cushman, Under fir and hemlock Olympic Mountains C. immixtus Kauffman 1932 20.10.1915 C.H. Kauffman KF732323 C. inamoenus (J. Favre) Quadr. 1985 (holotype)* 13522 G Switzerland C. rosargutus Chevassut & Rob. Henry 1978 Unknown J. Favre KF732324 In subalpine coniferous forest on calcareous ground C. infractus (Pers.: Fr.) Fr. 1838 (neotype) CFP495 S Sweden, Boh, Tossene, Anneröd In beech forest, on medium rich ground C. infractus (Pers.: Fr.) Fr. 1838 15.09.1986 T.E. Brandrud et al. KF732325 C. infractus var. aeruginosus Rob. Henry ex Reumaux 2009 XC2006-66 PC France, Seine-et-Marne, forêt de Villefermoy In deciduous forest on clayey-calcareous soil C. infractus (Pers.: Fr.) Fr. 1838 11.10.2006 R. Chalange KF732326 C. infractus var. flavus M.M. Moser 1999 (holotype)* 97/169 IB USA, Wyoming, Shoshone National Forest, Brooks Lodge In coniferous forest under Picea engelmannii and Abies lasiocarpa C. infractiflavus (M.M. Moser) Kytöv., 12.08.1997 Niskanen, Liimat., Bojantchev & Ammirati stat. nov. & nom. nov. M. Moser KF732327 C. josephii Reumaux 2006 (holotype) PML5193 PC France, Ardennes, forêt d’Elan In calcareous deciduous forest C. gracilior (Jul. Schäff. ex M.M. Moser) M.M. Moser 1967 P. Reumaux KF732328 08.10.1999 RH3880 PC France, Haut-Doubs, L’Hôpital du Grosbois In submontane, calcareous Carpinus forest C. juxtadibaphus Rob. Henry 1983 1950 Unknown KF732329 1965/0042 IB Austria, Tyrol, Oetztal, Untergurgl Under Alnus viridis C. kuehneri M.M. Moser 1974 03.09.1965 M. Moser KF732330 C. kytoevuorii Niskanen & Liimat. sp. nov. (holotype) TN05-158, H6029355 H Finland, Ks, Kuusamo, Oulanka, Ampumavaara In old, grass-herb Picea abies forest with some C. kytoevuorii Niskanen & Liimat. sp. nov. 17.09.2005 Betula, Pinus sylvestris and Populus tremula, on calcareous ground K. Liimatainen & T. Niskanen KF732529 C. laetargutus Chevassut & Rob. Henry 1978 (holotype) RH70405 PC France, Mont Aigoual In mixed forest of Picea and Fagus C. crassus Fr. 1838 29.09.1975 Mme Moutet KF732331 C. largoides Rob. Henry ex Bidaud, Carteret & Reumaux 2009 (holotype) PML2336 PC France, Ain, bois de la Morgne, Ordonnaz In deciduous forest C. subpurpurascens (Batsch) Fr. 1838 06.10.1991 A. Bidaud KF732332 C. largus Fr. 1838 (neotype) TN08-060 (H6001957) H Finland, V, Turku, Ruissalo In deciduous forest of Quercus robur, Corylus avellana and some Betula on mull soil C. largus Fr. 1838 03.09.2008 K. Liimatainen & T. Niskanen AB859985 C. largusiellus Reumaux 1996 (holotype) 3411 G France, Ardennes, Forêt de Boult Under deciduous trees (Betula, Carpinus) on clayey-calcareous soil C. largus Fr. 1838 09.10.1992 P. Reumaux KF732334 C. latoclaricolor Rob. Henry 1989 (holotype) RH1199 PC France In Picea forest C. badiolatus (M.M. Moser) M.M. Moser 1967 Unknown Unknown KF732335 103 C. juxtadibaphus Rob. Henry 1983 (holotype) C. kuehneri M.M. Moser 1974 (holotype) K. Liimatainen et al.: Identification and nomenclature of Phlegmacium C. fuligineofolius (M.M. Moser) M.M. Moser & Peintner 2002 (holotype) 104 Table 1 (cont.). Species Voucher C. lemanicus A. Favre & Vialard 2000 (holotype) 452177 C. leonicolor Reumaux 2001 (holotype) 4739 Herb. Locality Ecology Current name Collection date Collector GenBank number G France, Savoy, Thonon-les-Bains, Margencel Under Pinus and Picea on calcareous soil C. delaportei Rob. Henry 1988 06.11.1999 A. Favre, J. Vialard KF732336 PC France, Ardennes, bois du Vivier In clayey-calcareous deciduous forest under Quercus and Carpinus C. leonicolor Reumaux 2001 04.10.1992 P. Reumaux KF732337 C. lilacinicolor Reumaux 1996 (holotype) 3512 G France, Ardennes, La Croix-aux-Bois In deciduous forest on clayey-calcareous soil C. largus Fr. 1838 16.09.1979 P. Reumaux KF732338 C. lintrisporus Reumaux 1997 (holotype) 2935 G France, Ardennes, Mont Dieu In clayey-calcareous deciduous forest (Carpinus, Betula) C. largus Fr. 1838 23.09.1992 P. Reumaux KF732339 C. lividoviolaceus Rob. Henry 1987 (paratype)* RH2932 PC France In frondose woodland C. largus Fr. 1838 1969 Unknown KF732340 C. luteiaureus Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK07-247b (H6033617) H Finland, OP, Kiiminki, Juuvansydänmaa In grass-herb Picea abies forest with some Betula, Populus tremula and Pinus, on calcareous ground C. luteiaureus Kytöv., Liimat. & Niskanen sp. nov. 17.08.2007 I. Kytövuori KF732568 C. luteoarmillatus A.H. Sm. 1942 (holotype)* 15360 MICH USA, Michigan, Sharron Hollow On muck soil in low woods C. luteoarmillatus A.H. Sm. 1942 14.09.1940 A.H. Smith KF732341 C. luteobrunnescens A.H. Sm. 1944 (holotype) 17785 MICH USA, Washington, Olympic National Park, Olympic Hot Springs Under conifers C. luteobrunnescens A.H. Sm. 1944 11.10.1941 D.E. Stunz & A.H. Smith KF732342 C. luteocingulatus Bidaud & Fillion 1992 (holotype)* AB91-10-260 G France, Haute-Savoie, Forêt de la Semine Under Quercus and Carpinus on clayey, slightly acid soil C. luteocingulatus Bidaud & Fillion 1992 10.1991 Unknown KF732343 C. luteovaginans Bidaud & Faurite-Gendron 2006 (holotype) AB03-11-73 PC France, Ardèche, Saint-Alban-surSampzon In deciduous forest under Quercus ilex and Q. humilis C. aurantiopallidus Bidaud 2006 05.11.2003 A. Faurite-Gendron KF732344 C. maculatipes Bidaud 1996 (holotype) 2845 G France, Savoie, Les Arcs In subalpine zone under Picea abies and Alnus viridis C. pseudonaevosus Rob. Henry 1957 13.08.1992 P.A. Moreau KF732345 C. maculatocaespitosus Bidaud 2009 (holotype) AB08-10-302 PC France, Ain, Cerin In calcareous deciduous forest C. maculatocaespitosus Bidaud 2009 11.10.2008 A. Bidaud KF732346 C. mahiquesii Vila, A. Ortega & Suár.-Sant. 2008 (holotype) GDA54298 GDA Spain, Catalonia, Perafita Under Cistus monspeliensis, on acid soil C. mahiquesii Vila, A. Ortega & Suár.-Sant. 2008 18.01.2008 J. Vila, X. Llimona FM202139 C. melleicarneus Kytöv., Liimat., Niskanen & Brandrud sp. nov. (holotype) IK01-053 H Estonia, Hiiumaa, Sarve, Soonlepa In deciduous forest (Corylus, Quercus) with some Pinus on calcareous ground C. melleicarneus Kytöv., Liimat., Niskanen & Brandrud sp. nov. 16.09.2001 I. Kytövuori KF732577 C. mendax Bidaud, Mahiques & Reumaux 2011 (holotype) AB07-10-162 PC France, Ain, col de Richemont, Chanay In mixed forest under Betula and Picea C. mendax Bidaud, Mahiques & Reumaux 2011 12.10.2007 E. Bidaud & R. Fillion KF732401 C. metarius Kauffman 1921 (holotype) MICH10374 MICH USA, Colorado, Grand, Leal Under spruce and fir C. metarius Kauffman 1921 29.08.1917 C.H. Kauffman KF732347 C. misermontii Chevassut & Rob. Henry 1986 (holotype) RH84.134 PC France, Montpellier Under Quercus ilex C. misermontii Chevassut & Rob. Henry 1986 12.1984 G. Chevassut KF732348 C. montanus Kauffman 1932 (holotype) MICH10377 MICH USA, Oregon, Clackamas, Mt Hood, near Welches Post Office In hemlock and cedar forest C. montanus Kauffman 1932 03.10.1922 C.H. Kauffman KF732349 C. multiformis Fr. 1838 (neotype) CFP445 S Sweden, Ång, Häggdånger, Sjö In spruce forest with blueberry C. multiformis Fr. 1838 21.08.1986 T.E. Brandrud et al. KF732350 C. multiformis var. caesiophyllus Moënne-Locc. 2006 (holotype) PML882 PC France, Savoie, Arith Under Picea on calcareous soil C. caesiolamellatus (Bidaud) Kytöv., Liimat., Niskanen, Brandrud, Frøslev & A.F.S. Taylor comb. nov. 03.06.1988 P. Moënne-Loccoz KF732351 3582 G France, Haute-Savoie, Quintal Under Picea on calcareous soil C. variecolor (Pers.: Fr.) Fr. 1838 11.10.1993 P. Moënne-Loccoz KF732352 17451 MICH USA, Washington, Olympic National Park, Olympic Hot Springs Under conifers, pine, Douglas fir and mountain hemlock C. occidentalis A.H. Sm. 1939 30.09.1941 A.H. Smith KF732353 C. myrtilliphilus Kytöv., Liimat., Niskanen & Brandrud sp. nov. (holotype) IK97-1469 (H6032751) H Finland, Kn, Suomussalmi, Suolijärvi In gently sloping, partly swampy, grassherb spruce forest with some Pinus, Betula, Populus, Alnus incana and Salix spp. C. myrtilliphilus Kytöv., Liimat., Niskanen & Brandrud sp. nov. 14.09.1997 I. Kytövuori KF732605 C. neotriumphans Bidaud, Moënne-Locc. & Reumaux 2000 (holotype) PML209 G France, Haute-Savoie, Semnoz Under Picea on calcareous soil C. neotriumphans Bidaud, MoënneLocc. & Reumaux 2000 18.09.1985 P. Moënne-Loccoz KF732354 C. norrlandicus Brandrud 1989 (isotype) CFP526 S Sweden, Ång, Häggdånger, Torrom High herbaceous spruce forest on rich ground C. norrlandicus Brandrud 22.09.1986 T.E. Brandrud DQ117928 C. obsoletus Kühner 1955 (syntype) Sa-52-66 (G00262069) G France, Haute-Savoie, Samoëns Under Fagus C. obsoletus Kühner 1955 22.09.1952 R. Kühner KF732628 C. obsoletus Kühner 1955 (syntype) Sa-52-91 (G00262070) G France, Haute-Savoie, Samoëns Under Fagus and Quercus among moss and grass C. obsoletus Kühner 1955 27.09.1952 R. Kühner KF732356 C. occidentalis A.H. Sm. 1939 (holotype) 8654 MICH USA, California, Trinidad Under redwood C. occidentalis A.H. Sm. 1939 12.11.1937 A.H. Smith KF732357 Persoonia – Volume 33, 2014 C. muricinicolor Moënne-Locc. 1996 (holotype) C. mutabilis A.H. Sm. 1944 (holotype) 3591 G France, Ile-de-France, région de Versailles In calcareous deciduous forest C. largus Fr. 1838 C. ochraceobrunneus Rob. Henry ex Bidaud, Moënne-Locc. & Reumaux 2000 (holotype) 4027 G France, Haute-Savoie, Bois de Vorcier In mixed forest dominated by Picea C. ochribubalinus Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK93-641, H6032734 H Finland, U, Espoo, Nuuksio C. ochroclarus Rob. Henry ex Rob. Henry 1996 (holotype)* 2792 G C. ochropudorinus Rob. Henry ex Bidaud & Reumaux 2006 (holotype) PML2339 C. olidoamarus var. valentinus Mahiques & A. Favre 1999 (holotype) M. Cerutti KF732358 C. ochraceobrunneus Rob. Henry ex 05.11.1995 Bidaud, Moënne-Locc. & Reumaux 2000 H. Debauvais KF732359 In fairly rich grass-herb forest with Populus tremula, Betula, Alnus incana, Quercus, Corylus, Prunus padus, Salix spp., and some old pines and young spruces C. ochribubalinus Kytöv., Liimat. & Niskanen sp. nov. 02.09.1993 I. Kytövuori KF732530 France, Haute-Savoie, Forêt de la Semine In deciduous forest mixed with some Picea on clayey-calcareous soil C. ochroclarus Rob. Henry ex Rob. Henry 1996 11.07.1992 R. Fillion KF732360 PC France, Ain, Meyriat In deciduous forest C. talus Fr. 1838 09.10.1991 A. Bidaud KF732361 452173 G Spain, Valencia Under Quercus suber C. olidoamarus var. valentinus Mahiques & A. Favre 1999 26.10.1996 Unknown KF732362 C. oliveopetasatus M.M. Moser 2000 (holotype) 95/ 360 IB USA, Oregon, Wasco Co., Mt Hood, Clear Creek Camp Ground In mixed coniferous forest (Abies, Picea, Pinus, Tsuga, Pseudotsuga, Larix) C. oliveopetasatus M.M. Moser 2000 25.10.1995 M. Moser KF732363 C. olympianus A.H. Sm. 1939 (holotype) MICH USA, Washington, Olympic Mountains, Olympic Hot Springs, Boulder Creek Under fir C. olympianus A.H. Sm. 1939 19.10.1935 A.H. Smith KF732364 KF732365 3242 14.10.1993 C. ophiopus Peck 1878 (part of type)* s.n. PC USA, Maryland Among fallen leaves in woods C. ophiopus Peck 1878 September Unknown C. oregonensis A.H. Sm. 1939 (holotype) 3557 MICH USA, Oregon, Florence, Lake Tahkenitch Under spruce C. oregonensis A.H. Sm. 1939 19.11.1935 A.H. Smith KF732366 C. orichalceus var. olympianus A.H. Sm. 1944 (holotype) 17513 MICH USA, Washington, Olympic National Park, Olympic Hot Springs Under conifers C. pseudocupreorufus (A.H. Sm.) Nis02.10.1941 kanen, Liimat. & Ammirati stat. nov. & nom. nov. A.H. Smith KF732367 C. orichalceus var. olympianus f. luteifolius A.H. Sm. 1944 (holotype) 16970 MICH USA, Washington, Olympic Mts, Lake Angeles Under conifers C. luteicolor (A.H. Sm.) Ammirati, Bojant- 19.09.1941 chev, Niskanen & Liimat. stat. nov. & nom. nov. A.H. Smith KF732368 C. orichalceus var. xanthocephalus A.H. Sm. 1944 (holotype)* 17514 MICH USA, Washington, Olympic National Park, Olympic Hot Springs Under conifers C. orichalceus var. xanthocephalus A.H. Sm. 1944 02.10.1941 A.H. Smith KF732369 C. pallidifolius A.H. Sm. 1939 (holotype) 3244 MICH USA, Washington, Olympic Mountains, Olympic Hot Springs Under fir C. pallidifolius A.H. Sm. 1939 19.10.1935 A.H. Smith KF732370 C. pallidirimosus Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK95-585, H6035694 H Finland, InL, Utsjoki, Kevo, Tsieskuljohka In mesic heath forest with Betula and Pinus, some moist depressions C. pallidirimosus Kytöv., Liimat. & Niskanen sp. nov. 17.08.1995 I. Kytövuori KF732578 C. pansa (Fr.) Sacc. 1887 (neotype) IK90-1826 H Finland, V, Kemiö, Pederså At small, abandoned limestone quarries, spruce heath forest, roadside C. pansa (Fr.) Sacc. 1887 21.09.1990 I. Kytövuori KF732522 T.E. Brandrud et al. KF732371 C. papulosus Fr. 1838 (neotype)* CFP344 S Sweden, Mpd, Alnö, Ås In spruce forest on calcareous ground C. papulosus Fr. 1838 07.09.1985 C. paracrassus Reumaux 1995* 3522 G France, Ile-de-France, Fontainebleau In calcareous deciduous forest C. largus Fr. 1838 07.10.1979 N. Martelli KF732372 C. paracyanopus Moënne-Locc. & Reumaux 1996 (holotype) 3510 G France, Ardennes, Bois du Mont Dieu Under Carpinus and Betula, on clayeycalcareous soil C. largus Fr. 1838 25.09.1980 P. Reumaux KF732373 C. parafulmineus Rob. Henry 1993 2384 G France, Drôme, Romeyer In montane forest of Fagus and Abies, on calcareous ground C. parafulmineus Rob. Henry 1993 02.11.1991 A. Bidaud & P. Reumaux KF732552 C. parargutus Bidaud, Moënne-Locc. & Reumaux 1999 (holotype) 1144 G France, Ile-de-France In deciduous forest C. pardinus Reumaux 1995 24.08.1987 J. Poirier KF732374 C. pardinus Reumaux 1995 (holotype) 3432 G France, Ardennes, Forêt de Belval In deciduous forest on clayey-calcareous soil C. pardinus Reumaux 1995 27.09.1992 P. Reumaux & F. Reumaux KF732375 C. parolivascens Moënne-Locc. & Reumaux 2009 (holotype) PML29 PC France, Haute-Savoie, plateau des Glières Under Pinus among Sphagnum C. scaurus (Fr.: Fr.) Fr. 1838 08.09.1984 R. Baubet KF732377 C. patibilis Brandrud & Melot 1983 (holotype)* 213-78 O Norway, Akh, Nannestad, Hornsjøen In oligotrophic Picea abies forest C. patibilis Brandrud & Melot 1983 10.08.1978 T.E. Brandrud KF732378 C. patibilis var. scoticus Brandrud 1997 (holotype)* TEB161-83 E Scotland, Perthshire, Calvine, Struan Wood In subalpine zone under Betula pubescens C. largus Fr. 1838 22.09.1983 T.E. Brandrud KF732379 C. percomis Fr. 1838 (neotype) TN08-041 H Finland, V, Karjaa, Kohagen In herb-rich Picea abies forest with some Corylus avellana, Quercus robur, Betula and Populus tremula C. percomis Fr. 1838 02.09.2008 K. Liimatainen & T. Niskanen KF732380 C. perpallens Chevassut & Rob. Henry 1978 (holotype) RH3928 PC France, Mont Aigoual Under Picea and Fagus C. perpallens Chevassut & Rob. Henry 1978 28.10.1973 Unknown KF732381 AB97-11-496 PC France, Ain, Cerin In grassy meadow C. persoonianus Bidaud 2009 16.11.1997 A. Bidaud KF732382 RH2046 PC France, Jura, Forêt de Chaux In frondose forest with Quercus and Fagus C. crassus Fr. 1838 07.1965 R. Henry KF732383 C. pini Brandrud 1996 (isotype) CFP394 S Norway, Oppl, Østre Toten, Balke In coniferous forest on calcareous ground C. pini Brandrud 1996 25.09.1985 T.E. Brandrud et al. KF732384 C. piriodolens Moënne-Locc. 1996 (holotype) 642 G France, Haute-Savoie, Plateau de Solaison In young calcareous Picea forest C. variecolor (Pers.: Fr.) Fr. 1838 01.09.1987 R. Baubet KF732385 105 C. persoonianus Bidaud 2009 (holotype)* C. phylladus Rob. Henry 1983 (holotype)* K. Liimatainen et al.: Identification and nomenclature of Phlegmacium C. occultus Moënne-Locc. & Reumaux 1996 (holotype) 106 Table 1 (cont.). Species Voucher Herb. Locality Ecology Current name Collection date Collector GenBank number C. platypus (M.M. Moser) M.M. Moser 1967 (holotype) 58/64 (19580064) IB Germany, Baden-Württemberg, Inzigkofen In Fagus forest on calcareous soil C. platypus (M.M. Moser) M.M. Moser 1967 09.10.1948 M.M. Moser KF732563 KF732386 C. ponderosus A.H. Sm. 1939 (holotype) 9273 MICH USA, Oregon, Cave City Under Pinus ponderosa and Quercus spp. C. ponderosus A.H. Sm. 1939 01.12.1937 A.H. Smith C. porphyropus (Alb. & Schwein.) Fr. 1838 (neotype) CFP717 S Sweden, Jmt, Ragunda, Ragunda In birch forest on rich ground C. porphyropus (Alb. & Schwein.) Fr. 1838 20.08.1988 T.E. Brandrud et al. KF732387 C. porphyropus var. porphyrophorus Reumaux 2009 (holotype) PML5086 PC France, Ardennes, La Croix-aux-Bois In deciduous forest C. porphyropus (Alb. & Schwein.) Fr. 1838 06.10.1995 P. Reumaux KF732388 C. praestans (Cordier) Gillet 1874 (epitype) IK94-1861 H France, Ain, communen d’Echallan In Fagus forest with Picea C. praestans (Cordier) Gillet 1874 27.10.1994 I. Kytövuori KF732389 C. psalliotoides Chevassut & Rob. Henry 1978 (holotype)* RH3154 PC France, St. Mitre-les-Remparts Under Quercus ilex C. psalliotoides Chevassut & Rob. Henry 1978 22.11.1970 Unknown KF732390 C. pseudocephalixus Bidaud & Moënne-Locc. 2000 (holotype)* 4446 G France, Drôme, Forêt de Romeyer In mixed calcareous forest C. pseudocephalixus Bidaud & Moënne-Locc. 2000 02.11.1991 A. Bidaud KF732392 C. pseudocyanites var. paucus Reumaux 2005 (holotype) PML5261 PC France, Val-de-Marne, bois d’Ivry In deciduous forest C. cyanites Fr. 1838 17.10.1998 A. Bardet & R. Bardet KF732393 C. pseudogracilior Reumaux 2006 (holotype) PML4858 PC France, Dordogne, Tursac In calcareous deciduous forest C. pseudogracilior Reumaux 2006 26.10.1997 P. Reumaux KF732394 C. pseudolargus Rob. Henry 1987 (holotype) RH70218 PC France, Doubs In frondose and coniferous forest C. pseudonaevosus Rob. Henry 1957 Unknown Unknown KF732395 C. pseudominor Rob. Henry ex Reumaux 2006 (holotype) PML4750 PC France, Ardennes, Bois des Alleux In deciduous forest on clayey-calcareous soil C. talus Fr. 1838 27.09.1997 P. Reumaux KF732396 KF732397 C. pseudonaevosus Rob. Henry 1957 (type) RH162 PC France In montane Picea forest C. pseudonaevosus Rob. Henry 1957 Unknown Unknown C. pseudonebularis Moënne-Locc. 1996 (holotype) 42 G France, Haute-Savoie, Massif du Semnoz In subalpine Picea forest C. pseudonebularis Moënne-Locc. 1996 22.09.1985 P. Moënne-Loccoz KF732398 4401 G France, Ain, Le Poizat In coniferous forest on calcareous ground C. varius (Schaeff.: Fr.) Fr. 1838 14.10.1995 P.A. Moreau KF732399 4516 G France, Ain, Innimont In mixed forest C. varius (Schaeff.: Fr.) Fr. 1838 08.10.1990 D. Mazuir KF732400 C. pseudotalus Rob. Henry ex Bidaud & Reumaux 2006 (holotype) PML4859 PC France, Seine-Maritime, Forêt de la Londe-Rouvray Under deciduous trees on calcareous soil C. talus Fr. 1838 24.10.1997 J.-C. Malaval KF732402 C. pseudoturmalis Bidaud & Moënne-Locc. 2010 (holotype) PML3465 PC France, Haute-Savoie, Thorens-Glières In coniferous forest C. claricolor (Fr.) Fr. 1838 29.07.1993 A. Faurite-Gendron KF732403 C. pseudovariegatus M.M Moser 1999 (holotype) 97/ 296 IB USA, Wyoming, Shoshone National Forest, Lake east of Two Ocean Mt In subalpine forest under Picea engelmannii, Pinus albicaulius C. pseudovariegatus M.M Moser 1999 31.08.1997 M. Moser KF732404 C. pseudovarius Moënne-Locc. & Reumaux 2000 (holotype)* 4391 G France, Ile de France, Bois de Noisiel In calcareous deciduous forest C. luteocingulatus Bidaud & Fillion 1992 20.10.1996 G. Flantzer KF732405 C. purpurascens Fr. 1838 (neotype) IK98-2121 H Sweden, Nrk, Hidinge, Garphyttans NP In fairly rich spruce grass-herb forest with Corylus, Populus tremula, Betula and Quercus C. purpurascens Fr. 1838 20.09.1998 I. Kytövuori KF732406 C. rapaceoides Bidaud, G. Riousset & Riousset 2000 (holotype) AB97-12-527 G France, St. Rémy de Provence, Bouches du Rhône In deciduous forest C. rapaceoides Bidaud, G. Riousset & Riousset 2000 03.11.1997 G. Riousset & L. Riousset KF732407 C. reverendissimus Bidaud, Moënne-Locc. & Reumaux 2000 (holotype) 4667 G France, Haute-Loire, Forêt de Miaune In mixed calcareous forest C. reverendissimus Bidaud, MoënneLocc. & Reumaux 2000 16.10.1997 P. Chapon KF732408 C. rex­claricolorum Bidaud, Carteret & Reumaux 2010 (holotype) AB04-09-163 PC France, Ain, la Vèche, Chanay In coniferous forest C. rex­claricolorum Bidaud, Carteret & Reumaux 2010 15.09.2004 A. Bidaud & R. Fillion KF732409 C. rioussetiae Chevassut & Rob. Henry 1986 (holotype) RH84.70-78 PC France, Gravesen Under Populus alba C. rioussetiae Chevassut & Rob. Henry 1986 10.1984 Mme. Riousset KF732410 C. rosargutus Chevassut & Rob. Henry 1978 (holotype) RH70477 PC France, Haut-Doubs Under conifers C. rosargutus Chevassut & Rob. Henry 1978 Unknown Unknown KF732411 KF732412 C. rufior Reumaux 2000 (holotype) 1118 G France, Haute Ardenne In coniferous forest, under Picea C. varius (Schaeff.: Fr.) Fr. 1838 08.11.1988 G. Flantzer C. rufoallutus Rob. Henry ex Bidaud & Reumaux 2006 (holotype) PML635 PC France, Haute-Savoie, Plateau de Glières Under Picea C. rufoallutus Rob. Henry ex Bidaud & Reumaux 2006 26.08.1987 P. Moënne-Loccoz KF732413 & J. Melot C. rufoallutus var. caesiolamellatus Bidaud 2006 (holotype) PML4905 PC France, Ain, col des Bérentin Under Picea on calcareous soil C. caesiolamellatus (Bidaud) Kytöv., Liimat., Niskanen, Brandrud, Frøslev & A.F.S. Taylor comb. nov. 03.10.1993 A. Bidaud C. rufolatus Moënne-Locc. 1996 (holotype) 664 G France, Haute-Savoie, Plateau de Glières Under Picea abies C. rufolatus Moënne-Locc. 1996 26.09.1987 P. Moënne-Loccoz KF732415 KF732414 Persoonia – Volume 33, 2014 C. pseudopansa Bidaud 2000 (holotype) C. pseudopimus Rob. Henry ex Rob. Henry 2000 (holotype) CFP941 S Sweden, Ång, Säbrä, Överdal In dry spruce forest on rich ground C. russus Fr. 1838 03.08.1990 T.E. Brandrud et al. KF732416 2954 G France, Ile-de-France, Les Mureaux On river sand-banks under deciduous trees C. chromataphilus Rob. Henry 1989 01.10.1992 G. Redeuilh KF732417 C. saginoides Bidaud & Reumaux 2000 (holotype) 1264 G France, Ain, Forêt de Meyriat In mixed forest C. varius (Schaeff.: Fr.) Fr. 1838 22.10.1989 A. Bidaud & P. Reumaux KF732418 C. sannio M.M. Moser 1999 (holotype) 97/352 IB USA, Wyoming, Teton National Forest, Lost Lake In subalpine coniferous forest under Picea C. sannio M.M. Moser 1999 engelmannii, Abies lasiocarpa and Pinus albicaulis 10.09.1997 M. Moser KF732420 C. saxamontanus Fogel 1995 (holotype) F2535 MICH USA, Nevada, White Pine Co., Whealer Peak Campground Under Pinus spp. and Abies spp. C. saxamontanus Fogel 1995 30.06.1981 R. Fogel KF732421 C. scaurocaninus Chevassut & Rob. Henry 1982 (holotype)* RH71678 PC France, Montpellier Under Quercus ilex C. scaurocaninus Chevassut & Rob. Henry 1982 04.11.1979 G. Chevassut KF732422 C. scaurus (Fr.: Fr.) Fr. 1838 (neotype) CFP1074 S Switzerland, Bern, Fribourg, Rechthalten In the border of a peatbog with Pinus strobus C. scaurus (Fr.: Fr.) Fr. 1838 29.09.1991 T.E. Brandrud et al. KF732423 C. scaurus f. phaeophyllus M.M. Moser 2001 (holotype) 94/243 IB Sweden, Sm, Femsjö, Stora Mosse, Källanäset On bare peat soil in a mire with Pinus sylvestris C. scaurus (Fr.: Fr.) Fr. 1838 13.09.1994 M. Moser KF732424 C. scaurus subsp. violaceonitens Rob. Henry 1976 (holotype) RH2190 PC France In forest with Fagus, Quercus and Carpinus C. violaceonitens (Rob. Henry) Moënne-Locc. 2009 Unknown Unknown KF732425 C. scaurus var. notandus Bidaud 2009 (holotype) AB94-08-32 PC France, Loire, Jeansagnière In montane coniferous forest C. scaurus (Fr.: Fr.) Fr. 1838 21.08.1994 A. Bidaud KF732426 C. serariicolor Rob. Henry 1985 (holotype)* RH1731 PC France In montane coniferous forest C. papulosus Fr. 1838 Unknown Unknown KF732427 C. serarius Fr. 1838 (neotype) CFP959 S Sweden, Ång, Häggdånger, Torrom In dry spruce forest on rich ground C. serarius Fr. 1838 11.08.1990 Brandrud et al. KF732428 C. sobrius P. Karst. 1890 (type) PAK3235 H Finland, EH, Mustiala, Tammela In frondose forest C. sobrius P. Karst. 1890 23.09.1890 P.A. Karsten KF732429 C. sphagnetorum Bidaud 1996 (holotype) 3746 G France, Savoie, Lac du Clou - Beaufortin C. pseudonaevosus Rob. Henry 1957 31.08.1991 A. Bidaud KF732430 C. splendens var. papillatosporus Bidaud & Moënne-Locc. 2003 (holotype) 960 PC France, Haute-Savoie, Les Puisots In subalpine zone under Picea, among Sphagnum and Vaccinium In mixed forest C. splendens var. papillatosporus Bidaud & Moënne-Locc. 2003 25.09.1988 Exposotion of Annecy KF732431 C. squamosocephalus Bidaud, Moënne-Locc. & 99670 Reumaux 1999 (holotype) PC France, Ardennes, Bois de la Brouille Under Quercus C. squamosocephalus Bidaud, Moënne-Locc. & Reumaux 1999 02.10.1998 P. Reumaux KF732432 C. subaccedens Rob. Henry 1989 (holotype)* RH3170 PC France, Languedoc-RoussillonCévennes region Under Quercus ilex C. subaccedens Rob. Henry 1989 Unknown Unknown KF732433 C. subamaricatus Bidaud 2008 (holotype) AB94-10-313 PC France, Ain, Meyriat In calcareous mixed forest of Fagus and Abies C. tirolianus Bidaud, Moënne-Locc. & Reumaux 2005 16.10.1994 A. Bidaud KF732434 C. subaustralis A.H. Sm. & Hesler 1944 (holotype)* 14336 MICH USA, N.C., Great Smoky Mts National Park, Indian Gap In coniferous forest C. crassus Fr. 1838 14.07.1942 L.R. Hesler KF732435 C. subbalteatus Kühner 1955 (syntype)* 8-12 (G00262072) G France, Savoie, St-Bon, Praz In Picea forest C. balteatus (Fr.) Fr. 1838 10.09.1927 R. Kühner KF732437 C. subcrassoides Moënne-Locc. & Remaux 1995 (holotype) 363 G France, Savoie, Col des Saisies In herbaceous Picea abies forest C. pseudonaevosus Rob. Henry 1957 29.09.1986 R. Campia KF732438 C. subcrassus Rob. Henry 1983 (holotype)* RH71520 PC France, Doubs, Seloncourt Under conifers C. crassus Fr. 1838 30.09.1979 KF732439 KF732440 C. subcyanites Bidaud 2005 (holotype) PML5304 PC France, Ain, Le Poizat Under Picea C. cyanites Fr. 1838 05.09.1999 From exhibition of Seloncourt A. Bidaud C. subdecolorans M. Langl. & Reumaux 2000 (holotype) 4403 G France, Marne, Bazancourt In deciduous forest C. subdecolorans M. Langl. & Reumaux 2000 20.10.1991 M. Langlois KF732441 C. subdecoloratus Reumaux 2000 (holotype) 3951 G France, Ardennes, Bois des Alleux Under Betula C. ochraceobrunneus Rob. Henry ex 30.10.1994 Bidaud, Moënne-Locc. & Reumaux 2000 G. Laffond & P. Reumaux KF732442 C. subfoetens M.M. Moser & McKnight 1995 (holotype) 89/307 IB USA, Wyoming, Teton National Forest, Fourmile meadow Under Picea engelmannii C. subfoetens M.M. Moser & McKnight 1995 21.08.1989 K.H. McKnight KF732443 C. subfoetidus A.H. Sm. 1944 (holotype) 17778 MICH USA, Washington, Olympic National Park, Olympic Hot Springs Under conifers C. subfoetidus A.H. Sm. 1944 11.10.1941 A.H. Smith KF732444 C. subfraudulosus Kytöv., Liimat. & Niskanen sp. nov. (holotype) IK11-006 H Norway, Oppl, Lunner, Skøyenåsen In Picea abies forest with Corylus on calcareous ground C. subfraudulosus Kytöv., Liimat. & Niskanen sp. nov. 03.09.2011 I. Kytövuori KF732564 KF732445 AB97-10-339 PC France, Ain, Petaray, Aranc In calcareous coniferous forest C. subrugulosus Bidaud & Armada 2006 21.10.1997 A. Bidaud PML5119 PC France, Ardennes, bois de Toges In deciduous forest C. subpurpurascens (Batsch) Fr. 1838 06.10.1996 P. Reumaux KF732446 C. sublilacinopes Bidaud, Moënne-Locc. & Reumaux 2001 (holotype) PML4819 PC France, Seine-Maritime, La LondeRouvray In calcareous deciduous forest C. sublilacinopes Bidaud, Moënne-Locc. & Reumaux 2001 24.10.1997 L.C. Malaval KF732561 C. subolivascens A.H. Sm. 1944 (holotype) 14311 MICH USA, Washington, Olympic National Park, Deer Lake Under conifers C. subolivascens A.H. Sm. 1944 13.06.1939 A.H. Smith KF732447 107 C. subfuligineus Bidaud 2008 (holotype) C. subinops Reumaux 2009 (holotype) K. Liimatainen et al.: Identification and nomenclature of Phlegmacium C. russus Fr. 1838 (neotype) C. sabuletorum Redeuilh & Reumaux 1995 (holotype)* 108 Table 1 (cont.). Species Voucher Herb. Locality Ecology Current name Collection date Collector GenBank number C. subopimus Bidaud 1995 (holotype) 3477 G 05.08.1993 A. Bidaud KF732448 TN08-059 H In herbaceous, calcareous Picea abies forest near Alnus viridis In deciduous forest of Quercus robur, Corylus avellana and some Betula on mull soil C. balteatus (Fr.) Fr. 1838 C. subpurpurascens (Batsch) Fr. 1838 (epitype) France, Haute-Savoie, Plateau de Dran, Les Glières Finland, V, Turku C. subpurpurascens (Batsch) Fr. 1838 03.09.2008 K. Liimatainen & T. Niskanen KF732449 C. subpurpureophyllus A.H. Sm. 1939 (holotype) 8164 MICH USA, California, Crescent City Under spruce C. subpurpureophyllus A.H. Sm. 1939 28.10.1937 A.H. Smith KF732450 C. subrugulosus Bidaud & Armada 2006 (holotype) PC France, Isère, Treminis In coniferous forest on calcareous soil C. subrugulosus Bidaud & Armada 2006 06.10.2005 A. Bidaud & F. Armada KF732451 AB05-10-263 C. subsolitarius A.H. Sm. 1942 (holotype) 15377 MICH USA, Michigan, Ann Arbor On humus in oak-hickory woods C. subsolitarius A.H. Sm. 1942 11.09.1940 A.H. Smith KF732452 C. subspadiceus Reumaux 1996 (holotype) 2780 G France, Haute-Vienne, Limousin Under Picea abies C. largus Fr. 1838 15.10.1990 Taupenot KF732453 C. subtortus (Pers.: Fr.) Fr. 1838 (neotype) IK87-1510 S 18.09.1987 I. Kytövuori KF732454 4663 G In partly paludified spruce forest with some other tree species Under Quercus ilex on calcareous soil C. subtortus (Pers.: Fr.) Fr. 1838 C. subvariiformis Bidaud 2000 (holotype) Sweden, Sm, Hylte commune, Femsjö parish France, Bouches-du-Rhône, St Rémy de Provence C. luteocingulatus Bidaud & Fillion 1992 03.12.1997 G Riousset & L. Riousset KF732455 C. superbus A.H. Sm. 1944 (holotype) 17680 MICH USA, Washington, Olympic National Park, Olympic Hot Springs On steep mountain slopes C. superbus A.H. Sm. 1944 08.10.1941 A.H. Smith KF732456 C. talimultiformis Kytöv., Liimat., Niskanen, A.F.S. Taylor & Sesli sp. nov. (holotype) AT2004096 UPS Sweden, Upl, Hässelby Park In mixed forest C. talimultiformis Kytöv., Liimat., Niskanen, A.F.S. Taylor & Sesli sp. nov. 11.07.2004 A. Taylor KF732583 C. talus Fr. 1838 (neotype) CFP832 S Sweden, Jmt, Ragunda, Ragunda In birch forest on rich ground (Betula, Populus, Pinus) C. talus Fr. 1838 26.08.1989 T.E. Brandrud et al. KF732457 C. thalliopurpurascens Rob. Henry 1995 (isotype) RH458677 PC France, Jura, Bois Boucot In deciduous forest mainly of Carpinus and Fagus C. herpeticus Fr. 1838 10.1986 Unknown KF732458 C. tiliae Brandrud 1996 (holotype) TEB141-85 O Norway, Akh, Bærum, Løkkeåsen In dry calcareous soil, under Tilia cordata C. tiliae Brandrud 1996 28.08.1985 T.E. Brandrud KF732459 C. tirolianus Bidaud, Moënne-Locc. & Reumaux 2005 (holotype) PML4341 PC France, Jura, Moirans-en-Montagne Under Picea C. tirolianus Bidaud, Moënne-Locc. & Reumaux 2005 15.09.1996 A. Dégrange KF732460 C. triumphalis Bidaud, Moënne-Locc. & Reumaux 2000 (holotype) 3950 G France, Allier, Forêt des Colettes In coniferous forest C. triumphalis Bidaud, Moënne-Locc. & Reumaux 2000 23.10.1992 R. Chalange KF732462 C. turmalis Fr. 1838 (neotype) CFP716 S Sweden, Mpd, Borgsjö, Julåsen In herbaceous spruce forest C. turmalis Fr. 1838 20.08.1988 T.E. Brandrud et al. KF732464 C. vacciniophilus Brandrud 1997 (holotype) TEB17-88 O Norway, Oppl, Lunner, Søndre Oppdalen In forest of Picea abies C. pseudonaevosus Rob. Henry 1957 09.08.1988 T.E. Brandrud KF732465 C. van­campiae Cons. 2000* 871 MCVE Italy C. van­campiae Cons. 2000 04.10.1995 G. Consiglio JF907867 C. variecolor (Pers.: Fr.) Fr. 1838 (neotype) CFP1021 S Sweden, Gtl, Viklau, Tjakule In mixed woodland with Abies alba and Fagus sylvatica In spruce forest on calcareous ground C. variecolor (Pers.: Fr.) Fr. 1838 29.09.1990 T.E. Brandrud et al. KF732466 C. variipes Rob. Henry 1977 (holotype) RH5026 PC France, Ardennes In montane coniferous forest C. variipes Rob. Henry 1977 Unknown P. Reumaux KF732467 C. variosimilis M.M. Moser & Ammirati 1999 (holotype)* 89/493 IB USA, Washington, Skagit Co., Trail to Easy Pass In subalpine coniferous forest under Picea engelmannii, Abies lasiocarpa C. variosimilis M.M. Moser & Ammirati 1999 12.09.1989 M. Moser KF732468 CFP801 S Sweden, Ång, Häggdånger, Torrom In spruce forest on rich ground C. varius (Schaeff.: Fr.) Fr. 1838 23.09.1988 T.E. Brandrud et al. KF732469 MICH10435 MICH USA, Michigan, Washtenaw, Cascade Glen, Ann Arbor Among fallen leaves in mixed or frondose woods C. velicopius Kauffman 1918 29.09.1907 C.H. Kauffman KF732470 C. veneris Bidaud, Moënne-Locc. & Reumaux 1996 (holotype) 2952 G France, Haute-Loire, Dunières Under Abies alba on granitic soil C. flavescentipes Reumaux 1996 02.10.1992 B. Renon KF732471 C. violaceomaculatus Brandrud 1997 (holotype) CFP1449 S Sweden, Gtl, Bäl In forest of Picea abies and Pinus sylvestris on calcareous soil C. violaceomaculatus Brandrud 1997 28.09.1993 T.E. Brandrud et al. KF732473 C. violaceorubens Moënne-Locc. & Reumaux 1990 (holotype) PML005 G France, Haute-Savoie, Pessière plantée d’Avernioz Under Picea in needle litter C. violaceorubens Moënne-Locc. & Reumaux 1990 21.06.1985 P. Moënne-Loccoz KF732474 C. virentophyllus Kauffman 1918 (holotype) MICH10439 MICH USA, Michigan, Washtenaw, German Park Woods, SW of Ann Arbor On the ground, among grasses in frondose woods of oak, maple, etc. C. virentophyllus Kauffman 1918 11.10.1912 C.H. Kauffman KF732475 C. viridirubescens M.M. Moser & Ammirati 1997 (holotype) 95/ 688 IB USA, California, Mendocino Co., on Forest Under Lithocarpus densiflora and Road 408 about 8 miles from village Quercus garrayana C. viridirubescens M.M. Moser & Ammirati 1997 08.12.1995 M. Moser KF732476 C. vixolivascens Rob. Henry 1992 (holotype) RH89.123 PC France, Habsheim, Forêt de la Hardt C. vixolivascens Rob. Henry 1992 1989 M.V. Rastetter KF732477 In mixed deciduous forest with Carpinus, Acer, Crataegus and Quercus on subcalcareous soil C. volvatus A.H. Sm. 1939 (holotype) 8857 MICH USA, California, Crescent City Under spruce C. volvatus A.H. Sm. 1939 18.11.1937 A.H. Smith KF732478 C. wiebeae Thiers & A.H. Sm. 1969 (holotype) 8051 MICH USA, Oregon, Mt Hood, Camas Corral Under fir C. wiebeae Thiers & A.H. Sm. 1969 08.06.1958 E. Wiebe KF732479 Persoonia – Volume 33, 2014 C. varius (Schaeff.: Fr.) Fr. 1838 (neotype) C. velicopius Kauffman 1918 (lectotype)* K. Liimatainen et al.: Identification and nomenclature of Phlegmacium 109 Table 2 Cortinarius specimens other than types included in the phylogenetic analysis. Short sequences excluded from the analysis marked with *. Species Voucher Herb. Locality GenBank number C. acystidiosus Thiers 1960 C. aggregatus Kauffman 1918* C. alnobetulae Kühner 1989 C. alticaudus Reumaux 2008 C. areni­silvae (Brandrud) Brandrud 2012 C. argutus Fr. 1838 s. auct C. argutus s. auct. C. badiolatus (M.M. Moser) M.M. Moser 1967 C. balteatialutaceus Kytöv., Liimat. & Niskanen sp. nov. C. balteatialutaceus* C. balteatibulbosus Kytöv., Niskanen, Liimat., Bojantchev & A.F.S. Taylor sp. nov. C. balteatibulbosus* C. balteatibulbosus C. balteatibulbosus C. balteatibulbosus C. balteatibulbosus* C. balteatibulbosus C. balteatibulbosus C. balteatibulbosus C. balteatibulbosus* C. balteaticlavatus Kytöv., Liimat. & Niskanen sp. nov. C. balteaticlavatus C. balteaticlavatus C. balteatoalbus Rob. Henry 1985* C. balteatoalbus C. bigelowii Thiers & A.H. Sm. 1969 C. boreicyanites Kytöv., Liimat., Niskanen & A.F.S. Taylor sp. nov. C. boreicyanites C. boreidionysae Kytöv., Liimat., Niskanen & Dima sp. nov. C. borgsjoeensis Brandrud 1992 C. brunneiaurantius Kytöv., Liimat. & Niskanen sp. nov. C. brunneiaurantius C. caerulescens (Schaeff.) Fr. 1838 C. caesiocinctus C. caesiocolor Kytöv., Liimat. & Niskanen sp. nov. C. caesiolamellatus (Bidaud) Kytöv., Liimat., Niskanen, Brandrud, Frøslev & A.F.S. Taylor comb. nov. C. caesiolamellatus C. caesiolamellatus C. caesiophylloides Kytöv., Liimat., Niskanen, Brandrud & Frøslev sp. nov. C. caesiophylloides C. caesiophylloides C. caesiophylloides C. calojanthinus M.M. Moser & Ammirati 1999 C. calojanthinus C. castaneicolor A.H. Sm. 1944 C. chromataphilus Rob. Henry 1989 C. citriolens Ammirati & M.M. Moser 1999 C. claricolor (Fr.) Fr. 1838 C. claricolor C. cobaltinus Kytöv., Liimat. & Niskanen 2013 C. cobaltinus C. cremeiamarescens Kytöv., Liimat. & Niskanen sp. nov. C. cremeiamarescens C. cremeiamarescens C. cremeiamarescens C. cruentipellis Kytöv., Liimat., Niskanen & Dima sp. nov. C. cruentipellis C. cruentipellis C. cumatilis Fr. 1838 C. cumatilis C. cupreorufus Brandrud 1994 C. cupreorufus C. cupreorufus C. cyanites Fr. 1838 C. cyanites C. cyanites* C. cyanites C. dionysae Rob. Henry 1933 s. auct C. dionysae s. auct C. dionysae var. avellanus Rob. Henry ex Bidaud & Carteret 2008 C. eliae Bidaud, Moënne-Locc. & Reumaux 1996 C. elotoides M.M. Moser & McKnight 1995 C. evosmus Joachim ex Bidaud & Reumaux 2006 C. evosmus C. flavipallens Kytöv., Liimat. & Niskanen sp. nov. C. flavipallens C. aff. flavipallens C. flavobulbus Ammirati & M.M. Moser 1997 C. fraudulosus Britzelm. 1885 C. fraudulosus C. fuligineofolius (M.M. Moser) M.M. Moser & Peintner 2002 C. fuligineofolius C. gentianeus Bidaud 1993 C. gentianeus C. gentianeus C. gentianeus C. gentianeus CLO4681 TN10-179 JFA12247 IK09-1402b IK95-341 O-60164 T44 IK98-1029 JV10452, TUR5282 JR100900 DBB33060 TENN H WTU H H O O H TUR H UC USA Canada, QC, Papineauville Italy, Passo del Rolle Finland, PK, Kesälahti Finland, InL, Inari Norway Norway Sweden, Nb, Pajala Finland, InL, Utsjoki Norway, S&F, Sogndal Bulgaria, Saranzi KF732419 KF732512 EU655672 KF732610 KF732611 AY669535 UDB000138 KF732612 KF732587 KF732588 KF732590 DBB36892 H6027358 IK93-639, H6032755 IK04-044 H6032413 AT2004091 AT2004045 AT2004088 AT2004127 IK95-382, H6032729 IK96-782, H6032415 IK08-584, H6033533 CFP1083 IK97-761b OSC-81327 TN03-112 AT2010203 IK07-245 IK07-1029 IK93-644, H6032406 JV17979b, H6032422 UL98-88, TUB012146 IK97-1573 IK97-207, H6032730 TN09-201 UC H H H H UPS UPS UPS UPS H H H S H OSC H UPS H H H H TUB H H H Sweden, Järfälla Finland, V, Vihti Finland, U, Espoo Finland, U, Vantaa Finland, U, Helsinki Sweden, Upl, Berthåga graveyard Sweden, Upl, Slottsbacken Sweden, Upl, Stenhagen Sweden, Upl, Berthåga graveyard Finland, InL, Inari Finland, ES, Mäntyharju Finland, Ks, Taivalkoski Jurmu-Kurtti France, Ain, Ordonnaz Sweden, Jmt, Revsund USA, OR, Klamath Finland, PS, Kuopio Great Britain, Scotland, Grampian Finland, OP, Kiiminki Finland, Ks, Salla Finland, U, Espoo Finland, V, Turku Germany Finland Finland, U, Helsinki USA, WA, Olympic peninsula KF732591 KF732592 KF732593 KF732594 KF732595 UDB001132 UDB000711 UDB000715 UDB000722 KF732597 KF732598 KF732599 KF732613 KF732614 EU056976 KF732500 KF732501 KF732489 KF732615 KF732601 KF732602 AY174863 DQ663241 KF732604 KF732571 AT2005085 11841 IK92-3044 UPS TUB H Sweden, Upl Germany Finland, SoL, Pelkosenniemi UDB002202 AY669531 KF732573 IK08-1554 TF28 TF40 TN08-129 TSJ2003-005 SMI38 TUB 011862 19970154 TN07-138 TUB 011852 TSJ2006068 TF2006-103 IK00-027 TN06-273 OCS153F clone AlASOILE08 IK01-055 IK98-2503 IK11-008 IK92-2927 H6016455 TN02-700 TN07-378 TN11-129 IK98-1476 AT2000154 AT2004139 AT2004089 TUB011450 TSJ2000-102 IK94-1745a 41098 CFP503 TSJ2004-014 TUB011399 H6032393 TK368 SMIA06 JFA11826 TU106876 19960696 19910576 19910682 IK11-015 IK97-1378 IK09-1525 CFP775 TUB011845 H Finland, Kn, Paltamo H C UBC TUB IB H TUB C C H H UBC Finland, V, Västanfjärd Sweden, Vg Canada, BC Germany USA, WY Finland, Ks, Kuusamo Germany Norway, Oppl, Jevnaker Norway, Oppl, Jevnaker Finland, U, Siuntio Finland, V, Parainen Canada, BC USA, AK Estonia, Hiiumaa, Pühalepa Sweden, Öl, Algutsrum Norway, Akh, Asker Finland, V, Lohja Finland, U, Espoo Finland, Ks USA, WA, Olympic Peninsula USA, AK, Fairbanks Finland, U, Espoo Sweden, Upl, Uppsala Sweden, Upl, Uppsala Sweden, Upl, Uppsala Germany Germany, Bayern France, Ain, Oyonnax SE France Sweden, Vg Denmark Germany Finland, LK, Parikkala Finland, PeP, Tervola Canada, BC USA, CA, Del Norte Estonia, Saare, Kihelkonna Italy, Bozen KF732574 KF732576 KF732575 KF732538 DQ663281 FJ157126 AY669550 AF325607 KF732616 AY669522 KF673471 KF673472 KF732494 KF732495 EF218754 JN889840 KF732540 KF732541 KF732542 KF732517 KF732643 KF732548 KF732549 KF732550 KF732502 KF732503 KF732391 UDB001154 AY174813 DQ083782 KF732606 KF732617 DQ663395 DQ663368 AY174815 KF732555 KF732556 FJ039640 EU057017 UDB011906 UDB001036 AF478578 AF478577 KF732496 KF732497 KF732498 KF732499 AY669519 H H H H H H H H H UPS UPS UPS TUB C H PC S C TUB H H UBC WTU TU IB IB IB H H H S TUB Sweden, Gtl, Alskog and När parish Finland, Kn, Suomussalmi Finland, ES, Kerimäki Sweden, Dlr, Rättvik Germany 110 Persoonia – Volume 33, 2014 Table 2 (cont.) Species Voucher Herb. Locality GenBank number C. georgiolens Rob. Henry 1986 C. glaucocephalus M.M. Moser, Ammirati & Halling 2000 C. glaucopus (Schaeff.: Fr.) Gray 1821 C. glaucopus C. glaucopus C. glaucopus C. gracilior (Jul. Schäff. ex M.M. Moser) M.M. Moser 1967 C. herpeticus Fr. 1838 C. herpeticus C. herpeticus C. herpeticus C. immixtus Kauffman 1932 C. infractiflavus (M.M. Moser) Kytöv., Niskanen, Liimat., Bojantchev & Ammirati stat. nov. & nom. nov. C. infractiflavus C. infractiflavus C. infractus (Pers.: Fr.) Fr. 1838 C. infractus C. juxtadibaphus Rob. Henry 1983 C. largus Fr. 1838 C. leonicolor Reumaux 2001 C. luteicolor (A.H. Sm.) Ammirati, Bojantchev, Niskanen & Liimat. stat. nov. & nom. nov. C. luteicolor C. luteicolor C. luteicolor C. luteicolor C. luteobrunnescens A.H. Sm. 1944 C. maculatocaespitosus Bidaud 2009 C. meinhardii Bon 1986 s. auct. C. meinhardii s. auct. C. melleicarneus Kytöv., Liimat., Niskanen & Brandrud sp. nov. C. mendax Bidaud, Mahiques & Reumaux 2011 C. mendax C. metarius Kauffman 1921 C. misermontii Chevassut & Rob. Henry 1986 C. misermontii C. montanus Kauffman 1932 C. montanus C. multiformis Fr. 1838 C. multiformis C. multiformis C. multiformis C. multiformis C. multiformis C. multiformis C. multiformis C. multiformis C. multiformis C. myrtilliphilus Kytöv., Liimat., Niskanen & Brandrud sp. nov. C. neotriumphans Bidaud, Moënne-Locc. & Reumaux 2000 C. neotriumphans C. norrlandicus C. olivaceodionysae A. Ortega, Vila & Fdez.-Brime C. olivaceodionysae C. olivaceodionysae C. olivaceodionysae C. olivaceodionysae C. olivaceodionysae C. olivaceodionysae C. olivaceodionysae C. olivaceodionysae C. oliveopetasatus M.M. Moser 2000 C. olympianus A.H. Sm. 1939 C. ophiopus Peck 1878 C. ophiopus C. oregonensis A.H. Sm. 1939 C. orichalceus var. xanthocephalus A.H. Sm. 1944 C. orichalceus var. xanthocephalus C. orichalceus var. xanthocephalus C. palazonianus Vila, A. Ortega & Fdez.-Brime C. palazonianus C. pallidirimosus Kytöv., Liimat. & Niskanen sp. nov. C. pallidirimosus C. pallidirimosus C. pallidirimosus C. pallidirimosus C. pallidirimosus C. pansa (Fr.) Sacc. 1887 C. pansa C. pansa C. papulosus Fr. 1838 C. papulosus C. parafulmineus Rob. Henry 1993 C. pardinus Reumaux 1995 C. patibilis Brandrud & Melot 1983 C. patibilis C. patrickensis (M.M. Moser) Niskanen, Liimat., Kytöv., Bojantchev & Ammirati comb. nov.* C. patrickensis* IK98-2504 F19524 IK92-1105a SMIA20 clone NWBRO20 SMI293 JV19538 IK92-593 DB2142 TUB011456 9204 F17145OC73 DBB19634 H UBC H UBC UBC UC Sweden, Öl, Algutsrum Canada, BC Finland, SoL, Pelkosenniemi Canada, BC Canada, BC Canada, BC Italy, Veneto, Belluno Norway, Troms, Storfjord Hungary, Vas, Ispánk Germany Italy Canada, BC Bulgaria, Pirin Mts KF732618 HQ604682 KF732619 FJ039616 EU645632 FJ039615 KF732492 KF732507 KF732508 AY174808 JF907950 GQ159888 KF732534 IK92-1109 SMI286 TN02-832 IK93-223 CFP1108 CFP1085 CFP852 DBB46740 H UBC H H S S S UC Finland, SoL, Pelkosenniemi Canada, BC Finland, Ks, Oulanka Finland, EH, Pälkäne France, Ain France, Ain, Ordonnaz, Penant Belgium, Brabant, Tervuren USA, CA, Yosemite Nat’l Park KF732535 FJ039612 KF732523 KF732524 DQ663286 KF732333 KF732490 KF732546 DBB38211 JMB10-06-2007-03 VMS13 JFA11701 IK97-2298 TUB011396 TN05-168 TUB011443 O-125960 TN06-291 TN06-157 PML5204 IK98-2417 IK872172 OSC1064150 DBB00174 IK08-1857 TN06-139 IK98-1401 TN05-247 19940224 TAAM128778 TU105180 AT2004187 PK4471 F16414 O-125949 TN05-232 CFP475 IK99-711 TN06-311 IK94-1899 TN06-306 IK11-013 MK2540 IK12-001 IK07-1417 IK11-012 TUB011856 F14280 IK94-1225 IK01-050 AT2004256 F15822 CFP769 TN04-874 IK08-1482 TN04-1106 (H7017897) clone PS01-02 IK07-692 TN04-470 IK92-966 IK98-711 19990590 clone 8-73M8 IK97-1914 UC USA, CA, Yosemite Nat’l Park USA Canada, BC USA, OR, Clackamus Finland, V, Lohja UP21 TN06-319 IK90-1822 Arangu-Cort-03101201 RH70356 IK97-086 IK97-087, H6032748 DBB39406 IK95-1400, H7019584 UBC H, TUR H BP TUB PC H H UC KF732547 HM068562 FJ717511 EU057024 KF732620 AY174780 Finland, Ks, Kuusamo KF732551 Germany AY174840 Norway, AA, Grimstad AY669533 Finland, A, Sund KF732515 Finland, PK, Kitee KF732516 France, Ain DQ663236 Sweden, Öl, Högsrum KF732621 Spain, Catalonia, Girona KF732622 USA, OR EU525972 USA, OR JF795378 Finland, PS, Sonkajärvi KF732623 Finland, PK, Kitee KF732624 Finland, EH Vilppula KF732625 Norway, Hord, Voss KF732626 Sweden, Sm UDB001004 Estonia, Tartu UDB016114 Estonia, Voru UDB016130 Sweden, Jmtl UDB002163 Canada, BC FJ039634 Canada, BC FJ039635 Norway, Oppl, Ringebu AY669518 Norway, Hord, Ulvik KF732607 Sweden, Ång, Säbrå KF732608 Finland, PK, Juuka KF732627 Finland, SF, A, Jomala KF732480 France, Ain KF732481 Finland, A, Finström KF732482 Sweden, Gtl KF732483 Finland, A, Lemland KF732484 Estonia, Hiiumaa, Raplamaa KF732485 Sweden, Gstr KF732486 Sweden, Gtl KF732487 Germany AY669523 Canada, BC FJ157042 Finland, ES, Kerimäki KF732553 Finland, U, Helsinki KF732609 Sweden, Upl, Norrtälje UDB000729 Canada, BC FJ157035 Sweden, Dlr, Rättvik KF732543 Finland, V, Lohja KF732544 Finland, Kn, Paltamo KF732545 Italy, Sardinia, Nuoro, Gavoi, Lago di Gusana KF732531 Spain FJ946938 Finland, PeP, Tervola KF732579 Finland, PeP, Rovaniemi KF732580 Finland, SoL, Sodankylä KF732581 Norway, Troms, Storfjord KF732582 Russia, Sakha UDB001073 USA, OR JQ393042 Finland, ES, Mäntyharju KF732521 UK AM084701 Sweden DQ179120 Finland, A, Jomala KF732629 Finland, V, Kemiö KF732630 Spain, Roncal Navarra EF014269 France KF732461 Finland, V, Karkkila KF732631 Finland, V, Karkkila KF732632 USA, CA, Humboldt KF732532 H Sweden, Mpd, Borgsjö WTU H TUB H TUB O H H PC H H OSC UC H H H H IB TAAM TU UPS UBC UBC O H S H H H H H H H H H TUB UBC H H UPS UBC S H H H H H H H IB H H H KF732533 K. Liimatainen et al.: Identification and nomenclature of Phlegmacium 111 Table 2 (cont.) Species Voucher Herb. Locality C. percomis Fr. 1838 C. percomis C. percomis C. pini Brandrud 1996 C. porphyropus (Alb. & Schwein.) Fr. 1838 C. porphyropus C. praestans (Cordier) Gillet 1874 C. praestans C. praestans C. pseudocephalixus Bidaud & Moënne-Locc. 2000 C. pseudocephalixus C. pseudonaevosus Rob. Henry 1957 C. purpurascens Fr. 1838 C. purpurascens C. purpurascens C. purpurascens C. rapaceoides Bidaud, G. Riousset & Riousset 2000 C. cf. rhizophorus Bidaud & Cons. 2012 C. cf. rhizophorus C. cf. rhizophorus C. cf. rhizophorus C. rosargutus Chevassut & Rob. Henry 1978 CFP1104 AT2004243 TU105232 IK90-2288 TN10-004 TN06-151 TAAM128528 CFP482 TUB011460 IK98-1842 TUB011444 CFP1175 IK87-1174 IK09-1510 TUB011401 TAAM128795 TUB012692 IK95-1973 IK98-2451 TUB011860 JV01-574 IK96-1279 S UPS TU H H H TAAM S TUB H TUB S H H TUB TAAM TUB H H TUB C H France, Ain, Brenod Sweden, Upl Sweden, Gtl Finland, V, Parainen Canada, QC, Riviere-á-Pierre Finland, PK, Kitee Estonia, Saare Sweden, Upl, Börstil, Marieberg Germany Sweden, Ög, V. Tolstad Germany C. rosargutus C. rosargutus C. rufoallutus Rob. Henry ex Bidaud & Reumaux 2006 C. rufoallutus C. rufolatus Moënne-Locc. 1996 C. russus Fr. 1838 C. sannio M.M. Moser 1999 C. sannio C. saxamontanus Fogel 1995 C. scaurocaninus Chevassut & Rob. Henry 1982 C. scaurus (Fr.: Fr.) Fr. 1838 C. scaurus C. scaurus C. scaurus C. scaurus C. scaurus C. scaurus C. serarius Fr. 1838 C. sobrius P. Karst. 1890 Cortinarius sp. Cortinarius sp. Cortinarius sp.* C. spectabilis M.M. Moser 1952 s. auct. C. spectabilis s. auct. C. subdecolorans M. Langl. & Reumaux 2000 C. subfoetens M.M. Moser & McKnight 1995 C. subfoetens C. subfraudulosus Kytöv., Liimat. & Niskanen sp. nov. C. subfraudulosus C. subfraudulosus C. subfraudulosus C. sublilacinopes Bidaud, Moënne-Locc. & Reumaux 2001 C. subolivascens A.H. Sm. 1944 C. subpurpurascens (Batsch) Fr. 1838 C. subpurpurascens C. subpurpureophyllus A.H. Sm. 1939 C. subpurpureophyllus C. subpurpureophyllus C. subrubrovelatus (Bidaud) Kytöv., Liimat., Niskanen & Dima comb. nov. C. subrubrovelatus C. subrugulosus Bidaud & Armada 2006 C. subtortus (Pers.: Fr.) Fr. 1838 C. subtortus C. subtortus C. sulphurinus Quél. 1883 s. auct. C. sulphurinus var. fageticola Brandrud 1998 C. superbus A.H. Sm. 1944 C. talimultiformis Kytöv., Liimat., Niskanen, A.F.S. Taylor & Sesli sp. nov. C. talimultiformis C. talimultiformis C. talimultiformis C. tiliae Brandrud 1996 C. triumphalis Bidaud, Moënne-Locc. & Reumaux 2000 C. turmalis Fr. 1838 C. variegatus Bres. 1884 s. auct. C. variegatus s. auct. C. variipes Rob. Henry 1977 C. variosimilis M.M. Moser & Ammirati 1999 C. violaceonitens (Rob. Henry) Moënne-Locc. 2009 C. violaceonitens C. violaceorubens Moënne-Locc. & Reumaux 1990 C. violaceorubens C. viridirubescens M.M. Moser & Ammirati 1997 C. wiebeae Thiers & A.H. Sm. 1969 Hebeloma fastibile (Pers.) P. Kumm. H. mesophaeum (Pers.) Quél. IK07-242 IK08-565 AV010997 DBB00242 CFP1112 CFP923 IK98-891 IK88-1160 MTS-97-166-11 HS031095 TN10-053 TN03-1698 19940243 156946 19980175 19960092 AF-D35 TN04-923 IK04-045 IK03-005 IK03-004 IK11-010 TEB594-04 TEB595-04 IK11-011 IK08-2010 F17229OC157 CFP481 IK98-2245 IK88-2016 TU106607 TSJ2002-043 C1-EC172 AT2004275 14615 TN04-855 AT2005152 19890242 TUB011414 TU106663 IK98-2578 TN05-021 19760289 JMB07-08-2007-05 CFP506 CFP783 SMI45 IK300866 (H6032747) SES2741 TUB0118410 TAAM128693 O-63407 CFP781 IK92-1133 CFP525 F16442 CFP981 VMS26 TN06-170 TN00-661 TN07-062 TUB011885 JFA11817 clone NHPY20 19940036 GLM31004 H H S S H H WTU H H H IB TRTC IB IB H H H H H O O H H UBC S H H TU C WTU UPS H UPS IB TUB TU H H IB S S H TUB TAAM O S H S UBC S UBC H H H TUB WTU IB GenBank number KF732520 UDB000726 UDB015914 KF732633 KF732513 KF732514 UDB015946 KF732267 AY174804 KF732634 AY174784 KF732635 Finland, ES, Joutseno KF732511 Finland, LK, Parikkala KF732644 Germany AY174858 Estonia, Lääne-Viru UDB016117 Italy, Laina EU057049 Germany, Baden-Württemberg, Freudenstadt KF732569 Sweden, Öl, Borgholm KF732570 Germany AY669527 Denmark, Jylland DQ083813 Germany, Baden-Württemberg KF732636 Freiburg, Biederbach Finland, OP, Kiiminki KF732637 Finland, Ks, Taivalkoski KF732638 Sweden, Jmt (Borgsjö Congress) KF732639 USA, OR JF750423 France, Ain, Brenod KF732640 Sweden, Ång, Säbrå KF732519 Sweden, Nb, Pajala KF732536 Finland, KiL, Kittilä KF732537 USA, WA, Kittitas EU057026 Germany, Baden-Württemberg Karlsruhe KF732641 Canada, QC KF732509 Slovakia, Liptovská kotlina basin, Važec KF732510 Sweden, Femsjö UDB001068 Canada, ON JN021010 Sweden AF325562 Italy, Sudtirol UDB001069 Great Britain, Scotland, Banffshire UDB002441 Finland, U, Kirkkonummi KF732558 Finland, U, Helsinki KF732559 Sweden KF732525 Sweden, Öl, Nötbrunnskärret KF732526 Norway, Busk, Röyken KF732527 Norway, Oppl DQ663425 Norway, Oppl DQ663426 Norway KF732491 Finland, V, Lohja KF732560 Canada, BC GQ159817 Sweden, Upl, Börstils sn KF732565 Sweden, Srm, Mörkö parish KF732566 Sweden, Ög, Väversunda parish KF732567 Estonia, Saare, Kihelkonna UDB011261 Czech Republic DQ663434 USA, WA AY356323 Sweden, Upl UDB000736 Italy JF907905 Finland, U, Vantaa KF732557 Sweden, Upl UDB002241 USA, Wyoming, Yellowstone National Park GU363492 Germany AY174787 Estonia, Saare UDB011262 Sweden, Öl, Böda KF732463 Finland, ES, Joutsa KF732645 Sweden, Femsjö UDB001087 Canada, BC FJ717556 Sweden, Vg, Medelplana DQ663437 Sweden, Sk, Degeberga DQ663439 Canada, BC FJ157114 Finland, PH, Virrat KF732584 Turkey, Trabzon, Macka KF732585 Germany AY669532 Estonia, Tartu UDB015959 Norway AY669556 Sweden, Sk, Degeberga KF732642 Finland, SoL, Pelkosenniemi KF732436 Sweden, Upl, Vattholma KF732376 Canada, BC FJ039663 Sweden, Ång, Häggdånger KF732472 Canada, BC FJ717596 Finland, PK, Kitee KF732505 Finland, V, Kisko KF732506 Finland, V, Kisko KF732504 Germany AY669647 USA, CA, Mendocino EU057007 USA, OR/CA FJ440879 AF325643 AF126100 112 Persoonia – Volume 33, 2014 Hebeloma fastibile AF325643 Hebeloma mesophaeum AF126100 Cortinarius georgiolens IK98-2504 Sweden Cortinarius georgiolens TYPE France Cortinarius ponderosus AHS9273 TYPE U.S.A. Oregon Cortinarius perpallens RH3928 TYPE France 1.00 Cortinarius subtortus IK87-1510 TYPE Sweden Cortinarius subtortus FJ717556 Canada, British Columbia 0.88 0.01 substitutions/site 1.00 Cortinarius variipes RH5026 TYPE France Cortinarius vespertinus CFP981 Sweden 0.62 Cortinarius variipes (= C. vespertinus s. auct.) 1.00 Cortinarius oregonensis AHS3557 TYPE U.S.A. Oregon Cortinarius oregonensis FJ157035 Canada, British Columbia Cortinarius largoides PML2336 TYPE France Cortinarius subpurpurascens UDB000736 Sweden 1.00 Cortinarius subpurpurascens Cortinarius subinops PML5119 TYPE France Cortinarius subpurpurascens TN08-059 TYPE Finland 0.93 Cortinarius subpurpurascens JF907905 Italy 0.67 Cortinarius collocandoides PML5087 TYPE France Cortinarius collocandoides Cortinarius genuinus XC2005-132 TYPE France Cortinarius mendax TN06-291 Finland Cortinarius mendaz 1.00 Cortinarius mendax TN06-157 Finland (= C. subporphyropus s. auct.) Cortinarius mendaz AB07-10-162 TYPE France 0.88 Cortinarius purpurascens IK87-1174 Finland Cortinarius purpurascens IK09-1510 Finland 1.00 Cortinarius purpurascens Cortinarius eumarginatus AB07-10-175 TYPE France /Purpurascentes Cortinarius purpurascens IK98-2121 TYPE Sweden Cortinarius purpurascens UDB016117 Estonia Cortinarius porphyropus TN06-151 Finland Cortinarius porphyropus CFP717 TYPE Sweden 0.98 Cortinarius porphyropus Cortinarius porphyropus var. porphyrophorus PML5086 TYPE France Cortinarius porphyropus TN10-004 Canada, Québec Cortinarius mutabilis AHS17451 TYPE U.S.A. Washington 1.00 Cortinarius occidentalis Cortinarius occidentalis AHS8654 TYPE U.S.A. California Cortinarius amnicola AHS15381 TYPE U.S.A. Michigan Cortinarius cacodes IB91/618 TYPE U.S.A. California Cortinarius citrinipedes AHS15305 TYPE U.S.A. Michigan 0.68 Cortinarius rapaceoides 1.00 Cortinarius rapaceoides AB9712527 TYPE France 0.73 (= C. caroviolaceus s. auct.) Cortinarius caroviolaceus EU057049 Germany Cortinarius evosmus PML4837 TYPE France 1.00 Cortinarius evosmus DQ663368 Denmark (= C. osmophorus s. auct.) Cortinarius evosmus AY174815 Germany Cortinarius sannio IB97/352 TYPE U.S.A. Wyoming 1.00 Cortinarius sannio IK88-1160 Finland Cortinarius sannio IK98-891 Sweden Cortinarius arenicola AHS15315 TYPE U.S.A. Michigan Cortinarius pseudogracilior PML4858 TYPE France 1.00 Cortinarius sublilacinopes PML4819 TYPE France Cortinarius sublilacinopes DQ663434 Czech Republic 1.00 Cortinarius metarius MICH10374 TYPE U.S.A. Colorado Cortinarius metarius Cortinarius barbarorum DQ663236 TYPE France 1.00 Cortinarius frondosophilus PML4817 TYPE France 0.71 Cortinarius platypus Cortinarius platypus IB58/64 TYPE Germany Cortinarius spectabilis DQ663425 Norway 1.00 0.78 Cortinarius spectabilis DQ663426 Norway Cortinarius cf. flavipallens FJ039640 Canada, British Columbia 0.99 Cortinarius flavipallens IK08-1729 TYPE Finland /Calochroi p.p. 1.00 1.00 Cortinarius flavipallens H6032393 Finland Cortinarius flavipallens TK368 Finland Cortinarius calojanthinus IB97/220 TYPE U.S.A. Wyoming 1.00 Cortinarius calojanthinus Cortinarius corrosus TN08-129 Finland (= C. corrosus s. auct.) 0.89 Cortinarius corrosus DQ663281 Sweden Cortinarius cobaltinus KF673470 TYPE Finland 1.00 Cortinarius cobaltinus KF673471 Norway 0.96 1.00 Cortinarius cobaltinus KF673472 Norway 0.99 Cortinarius caesiocinctus DQ663239 TYPE France Cortinarius caesiocinctus DQ663241 Finland 1.00 Cortinarius olympianus AHS3242 TYPE U.S.A. Washington Cortinarius olympianus IK94-1225 Finland 0.64 1.00 Cortinarius bigelowii 45385 TYPE U.S.A. Idaho Cortinarius bigelowii EU056976 U.S.A. Oregon 1.00 Cortinarius elotoides IB87/60 TYPE U.S.A. Wyoming Cortinarius elotoides (= C. pseudoglaucopus s. auct.) Cortinarius pseudoglaucopus DQ663395 Sweden Cortinarius luteicolor FJ717511Canada British Columbia C. orich. var. olymp. f. luteifolius AHS16970 TYPE U.S.A. Washington 1.00 Cortinarius luteicolor HM068562 U.S.A. C. luteicolor Cortinarius luteicolor DBB46740 U.S.A. California Cortinarius luteicolor DBB38211 U.S.A. California Cortinarius luteicolor EU057024 U.S.A. Oregon 1.00 0.99 0.94 0.98 1.00 1.00 C. orichalceus var. zanthocephalus 1.00 Cortinarius aureofulvus CFP769 Sweden Cortinarius aureofulvus TN04-874 Finland AHS17514 TYPE U.S.A. Washington (= C. aureofulvus s. auct.) 0.62 Cortinarius aureofulvus IK08-1482 Finland Cortinarius sazamontanus F2535 TYPE U.S.A. Nevada Cortinarius sazamontanus EU057026 U.S.A. Washington C. orich. var. olympianus AHS17513 TYPE U.S.A. Washington C. pseudocupreorufus Cortinarius cupreorufus TN11-129 U.S.A. Alaska 1.00 Cortinarius cupreorufus TN07-378 U.S.A. Washington 0.92 Cortinarius cupreorufus TN02-700 Finland Cortinarius cupreorufus CFP1038 TYPE Sweden /Calochroi & Fulvi /Laeticolores Fig. 1 The Bayesian 50 % majority-rule consensus tree inferred from ITS regions. PP > 0.50 are indicated above branches. In blue colour and bold are new species described in this study as well as new combinations made and in orange and bold neo- and epitypes designated in this study. In pink are species described by French authors, in red species described by other European authors, in green species described by North American authors and in black other specimens included in this study. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium 0.64 0.85 0.64 1.00 Cortinarius sulphurinus DQ663437 Sweden Cortinarius sulphurinus s. auct. Cortinarius sulphurinus var. fageticola DQ663439 TYPE Sweden 1.00 Cortinarius oliveopetasatus IB95/360 TYPE U.S.A. Oregon 0.95 Cortinarius oliveopetasatus FJ157042 Canada, British Columbia 1.00 Cortinarius juxtadibaphus RH3880 TYPE France Cortinarius juxtadibaphus (= C. dibaphus s. auct.) Cortinarius dibaphus DQ663286 France 1.00 Cortinarius flavobulbus IB95/629 TYPE U.S.A. California Cortinarius flavobulbus EU057017 U.S.A. California Cortinarius subpurpureophyllus AHS8164 TYPE U.S.A. California Cortinarius subpurpureophyllus 1.00 Cortinarius napus TN04-855 Finland (= C. napus s. auct.) Cortinarius napus UDB002241 Sweden 113 /Calochroi & Fulvi Cortinarius napus GU363492 U.S.A. Wyoming 1.00 Cortinarius viridirubescens IB95/688 TYPE U.S.A. California Cortinarius viridirubescens EU057007 U.S.A. California Cortinarius alnobetulae 1.00 Cortinarius alnobetulae Kühner 53-6 TYPE France (= C. moseri s. auct.) Cortinarius alnobetulae EU655672 Italy 1.00 Cortinarius parafulmineus TYPE France Cortinarius parafulmineus EF014269 Spain Cortinarius meinhardii TN05-168 Finland 1.00 C. splendens var. papillatosporus PC960 TYPE France C. splendens var. papillatosporus (= Cortinarius meinhardii s. auct.) Cortinarius meinhardii AY174840 Germany 1.00 Cortinarius maculatocaespitosus AB08-10-302 TYPE France Cortinarius maculatocaespitosus AY174780 1.00 C. anfractoides var. cinereoclarus AB08-09-155 TYPE France C. persoonianus AB97-11-496 TYPE France Cortinarius infractiflavus FJ039612 Canada, British Columbia Cortinarius infractiflavus 1.00 1.00 Cortinarius infractiflavus IK92-1109 Finland (= C. infractus var. flavus IB97/169 TYPE 0.83 /Infracti U.S.A Wyoming) Cortinarius infractiflavus DBB19634 Bulgaria 0.97 C. infractus var. aeruginosus XC2006-66 TYPE France Cortinarius amarocaerulescens AB99-11-362 TYPE France 1.00 Cortinarius infractus Cortinarius infractus CFP495 TYPE Sweden Cortinarius infractus TN02-832 Finland Cortinarius infractus IK93-223 Finland 1.00 Cortinarius crassus CFP938 TYPE Sweden Cortinarius crassus Cortinarius laetargutus RH70405 TYPE France Cortinarius rufoallutus JF750423 U.S.A. Oregon 1.00 Cortinarius rufoallutus AV010997 Sweden 0.99 Cortinarius rufoallutus PML635 TYPE France Cortinarius melleicarneus IK01-053 TYPE Estonia 0.99 Cortinarius melleicarneus AY669533 Norway 0.73 Cortinarius talimultiformis IK300866 Finland C. aurantionapus var. similis PML883 TYPE France 0.98 Cortinarius talimultiformis SES2741 Turkey Cortinarius talimultiformis Cortinarius talimultiformis AY669532 Germany Cortinarius talimultiformis AT2004096 TYPE Sweden 1.00 Cortinarius talimultiformis UDB015959 Estonia Cortinarius multiformis FJ039635 Canada British Columbia Cortinarius multiformis FJ039634 Canada British Columbia 0.69 Cortinarius multiformis UDB002163 Sweden 0.90 1.00 Cortinarius multiformis TN05-247 Norway Cortinarius multiformis UDB016114 Estonia Cortinarius multiformis UDB016130 Estonia 0.94 Cortinarius multiformis IK08-1857 Finland Cortinarius multiformis TN06-139 Finland Cortinarius multiformis CFP445 TYPE Sweden /Multiformes 0.76 Cortinarius multiformis IK98-1401 Finland Cortinarius multiformis UDB001004 Sweden Cortinarius crenulatus PML4866 TYPE France 1.00 Cortinarius pseudotalus PML4859 TYPE France 1.00 Cortinarius talus Cortinarius pseudominor PML4750 TYPE France Cortinarius ochropudorinus PML2339 TYPE France Cortinarius talus CFP832 TYPE Sweden C. rufoallutus var. caesiolamellatus PML4905 TYPE France Cortinarius caesiolamellatus TN09-201 U.S.A. Washington 1.00 Cortinarius caesiolamellatus Cortinarius caesiolamellatus UDB002202 Sweden Cortinarius caesiolamellatus AY669531 Germany C. multiformis var. caesiophyllus PML882 TYPE France 1.00 Cortinarius caesiophylloides TN05-016 TYPE Finland 1.00 Cortinarius caesiophylloides TF2006-112 Cortinarius caesiophylloides IK92-3044 Finland 1.00 Cortinarius caesiophylloides IK08-1554 Finland Cortinarius pallidirimosus UDB001073 Russia 0.96 Cortinarius pallidirimosus JQ393042 U.S.A. Oregon 0.80 Cortinarius pallidirimosus IK92-966 Finland 0.92 Cortinarius pallidirimosus IK95-585 TYPE Finland 1.00 Cortinarius pallidirimosus TN04-470 Finland 1.00 Cortinarius pallidirimosus IK07-692 Finland 0.90 Cortinarius pallidirimosus IK98-711 Norway Cortinarius virentophyllus MICH10439 TYPE U.S.A. Michigan Cortinarius montanus MICH10377 TYPE U.S.A. Oregon 1.00 Cortinarius montanus EU525972 U.S.A. Oregon 0.98 Cortinarius montanus JF795378 U.S.A. Oregon /Scauri Cortinarius violaceonitens TN06-170 Finland 0.99 0.96 Cortinarius violaceonitens TN00-661 Finland Cortinarius violaceonitens RH2190 TYPE France (as C. scaurus subsp. violaceonitens) Fig. 1 (cont) 114 Persoonia – Volume 33, 2014 Cortinarius cd rpd tihu s IK92-593 Norway Cortinarius herpeticus CFP936 TYPE Sweden Cortinarius tc all iopurpurashd ns RH458677 France 0.55 Cortinarius cd rpd tihu s Cortinarius cd rpd tihu s DB2142 Hungary 0.61 Cortinarius cd rpd tihu s AY174808 Germany Cortinarius cd rpd tihu s JF907950 Italy 1.00 Cortinarius mul ie ind om ol ius AF478578 IB19910576 1.00 Cortinarius mul ie ind om ol ius IB91/682 TYPE U.S.A. California Cortinarius mul ie ind om ol ius AF478577 TYPE U.S.A. California /Scauri Cortinarius sh aurus TN03-1698 Slovakia 0.99 Cortinarius sh aurus JN021010 Canada Ontario Cortinarius sh aurus TN10-053 Canada Québec Cb sh aurus f. pc ad opccllu s IB94/243 TYPE Sweden 0.51 Cortinarius scaurus CFP1074 TYPE Switzerland 1.00 Cortinarius sh aurus Cb sh aurus f. pc ad opccllu s UDB001068 Sweden 0.59 Cortinarius sh aurus AF325562 Sweden Cortinarius sh aurus UDB001069 Italy Cortinarius sh aurus UDB002441 Scotland iv ashd ns PML29 TYPE France 0.90 Cortinarius parol Cm sh aurus var . notandus AB94-08-32 TYPE France aride atus s. auct. CFP525 Sweden 1.00 Cortinarius v Cortinarius v aride atus s. auct. FJ039663 Canada British Columbia Cortinarius h orrue is AHS16842 TYPE U.S.A. Washington 1.00 Cortinarius turf al is IK92-1133 Finland Cortinarius turf al is Cortinarius turmalis CFP716 TYPE Sweden Cortinarius sukm od tidus AHS17778 TYPE U.S.A. Washington Cortinarius v iol ahd orukd ns TN07-062 Finland 1.00 Cortinarius v iol ahd orukd ns PML005 TYPE France 0.95 Cortinarius v iol ahd orukd ns AY669647 Germany 0.52 Cortinarius boreicyanites CFP931 TYPE Sweden 1.00 Cortinarius k ord ihc anitd s TN03-112 Finland Cortinarius k ord ihc anitd s AT2010203 Scotland 1.00 / Cyanites Cortinarius sukhc anitd s PML5304 TYPE France Cortinarius hc anitd s IK98-1476 Finland sdu. ohc panitd s var. pauhu s PML5261 TYPE France 1.00 Cb Cortinarius hc anitd s Cortinarius hc anitd s AT2000154 Sweden Cortinarius hc anitd s UDB001154 Sweden Cortinarius cyanites AT2005069 TYPE Sweden Cortinarius v iol ahd of ahul atus CFP1449 TYPE Sweden Cortinarius ochribubalinus IK93-641 TYPE Finland Cortinarius hdpc al iw oidd s IB87/188 TYPE U.S.A. Wyoming 0.55 rku s AHS17680 TYPE U.S.A. Washington 1.00 Cortinarius supd Cortinarius supd rku s FJ157114 Canada British Columbia 0.67 0.90 Cortinarius alk om rae rans IB95/595 TYPE U.S.A. California Cortinarius v iw ol iv ashd ns RH89.123 TYPE France 0.71 ol oratus PML3951 TYPE France 1.00 Cortinarius suk.dh Cortinarius ohc rahd ok runndu s Cortinarius ohc rahd ok runndu s PML4027 TYPE France Cortinarius papul o sus TN06-319 Finland 1.00 0.97 Cb apul posus CFP344 TYPE Sweden 0.99 osus IK90-1822 Finland 0.98 Cortinarius papul 0.82 Cortinarius h astand ih ol or AHS17926 TYPE U.S.A. Washington 1.00 Cortinarius h astand ih ol or FJ157126 Canada British Columbia td ok runnd shd ns AHS17785 TYPE U.S.A. Washington 0.98 Cortinarius lu Cortinarius lu td ok runnd shd ns IK97-2298 Finland 0.94 / Elastici & Percomes Cortinarius luteiaureus IK07-247b TYPE Finland 1.00 Cortinarius serarius CFP959 TYPE Sweden Cortinarius sd rarius TN04-923 Finland 0.73 Cortinarius h itrinim ol ius AHS3158 TYPE U.S.A. Washington 1.00 Cortinarius pd rh of is CFP1104 France 1.00 Cortinarius percomis TN08-041 TYPE Finland Cortinarius pd rh of is UDB000726 Sweden 0.50 Cortinarius pd rh of is UDB015914 Sweden aportd i RH8673 TYPE France 1.00 Cortinarius ddl 0.53 Cortinarius ddl aportd i Cortinarius ldf anihu s G452177 TYPE France Cortinarius pall idim ol ius AHS3244 TYPE U.S.A. Washington ohdpc al iwu s IK98-1842 Sweden 1.00 Cortinarius psdu. Cortinarius psdu. ohdpc al iwu s 4446 TYPE Cortinarius h inh tipd s GE 02-100 TYPE France France Cortinarius psdu. ohdpc al iwu s AY174784 Germany Cortinarius ahc stidiosus CLO U.S.A. Tennessee Cb ahc stidiosus 1902 TYPE U.S.A. Texas IK95-1973 Germany Cortinarius cf. rc i 0.53 i IK98-2451 Sweden 1.00 Cortinarius cf. rc (= Cb iduhcu hls s. auct. p.p.) AY669527 Germany Cortinarius cf. rc i 1.00 / Elastici & Percomes s. lato Cortinarius cf. rc i DQ083813 Denmark td ov ae inans AB03-11-73 TYPE France 1.00 Cortinarius lu Cortinarius aurantiopall idus ish idoaf v arus ) Cortinarius aurantiopall idus AB05-11-404 TYPE France (= Cb 1.00 Cortinarius russus CFP941 TYPE Sweden Cortinarius russus CFP923 Sweden Cortinarius iff iw tus GQ159888 Canada British Columbia Cortinarius sae inus CFP475 Sweden 0.98 1.00 Cortinarius nd otriufpc ans Cortinarius sae inus TN05-232 Norway (= Cb sae inus s. auct.) / Phlegmacium 1.00 Cortinarius nd otriufpc ans G40 TYPE France andihu s DQ117928 TYPE Sweden 1.00 Cortinarius norrl Cortinarius norrl andihu s IK99-711 Finland sold tus 5a-52-66 SYNTYPE France 1.00 Cortinarius ok Cortinarius ok sold tus 5a-52-91 SYNTYPE France Fig. 1 (cont) 0.75 0.75 K. Liimatainen et al.: Identification and nomenclature of Phlegmacium 1.00 0.99 0.55 1.00 0.66 0.68 1.00 1.00 0.77 0.63 0.77 0.99 1.00 1.00 0.57 1.00 0.75 0.61 0.84 1.00 1.00 0.93 115 ah is 3950 TYPE France 1.00 Cortinarius triucdp Cortinarius triucdp ah is (= C. luhd inus s. auct., C. ophiopus s. Kühner) Cortinarius luhd inus CFP781 Sweden Cortinarius pseue ol arieb atus IB97/296 TYPE U.S.A. Wyoming 1.00 atih is H6016455 Finland 1.00 Cortinarius cuc Cortinarius cumatilis IK98-2164 TYPE Sweden Cortinarius praestans UDB015946 Estonia 1.00 Cortinarius praestans IK94-1861 TYPE France Cortinarius praestans AY174804 Germany /Claricolores Cortinarius rex.ch aric oh oruu AB04-09-163 TYPE France 1.00 Cortinarius pseue oturc ah is PML3465 TYPE France Cortinarius ch aric oh or TN07-138 Finland 0.96 Cortinarius ch aric oh or Cortinarius claricolor CFP691 TYPE Sweden Cortinarius ch aric oh or AY669522 Germany iae TEB141-85 TYPE Norway 1.00 Cortinarius tih Cortinarius tih iae AY669556 Norway Cortinarius fump neri IB1965-0042 TYPE Austria inus 3432 TYPE France 1.00 Cortinarius pare Cortinarius pare inus Cortinarius pararbu tus 1144 TYPE France Cortinarius patiw ih is IK97-086 Finland C. patiw ih is 213-78 TYPE Norway Cortinarius patiw ih is IK97-087 Finland orb sy oeensis IK07-1029 Finland 1.00 Cortinarius w Cortinarius w orb sy oeensis CFP728 TYPE Sweden Cortinarius c onbmc inus 3422 TYPE France Cortinarius ch al icmd s 2932 TYPE France Cortinarius ch arus 4038 TYPE France Cortinarius cud reol ioh acmu s 3426 TYPE France Cortinarius suw spae icmu s 2780 TYPE France tus 3591 TYPE France 1.00 Cortinarius occuh Cortinarius h arbu s Cortinarius paracy anopus 3510 TYPE France Cortinarius h intrisporus 2935 TYPE France Cortinarius h ih ac inic oh or 3512 TYPE France 0.97 Cortinarius largus TN08-060 TYPE Finland Cortinarius cmdp ah ixoh arbu s RH6048 TYPE France Cortinarius h arbu siehhu s 3411 TYPE France Cortinarius squac osocmdp ahu s PC99670 TYPE France 0.99 Cortinarius eh iae PML1032 TYPE France 1.00 Cortinarius eh iae PC41098 France (as C. cud reol ioh acmu s ) Cortinarius variecolor CFP1021 TYPE Sweden 1.00 Cortinarius l ariec oh or Cortinarius pirioe ohm ns 642 TYPE France Cortinarius cu ric inic oh or PML3582 TYPE France Cortinarius ruqoh atus PML664 TYPE France Cortinarius ruqoh atus (= C. spae icmhhu s s. auct.) Cortinarius spae icmhhu s CFP1112 France Cortinarius w ah teatich al atus IK95-382 Finland 1.00 Cortinarius balteaticlavatus IK96-595 TYPE Finland Cortinarius w ah teatich al atus IK96-782 Finland Cortinarius w ah teatich al atus IK08-584 Finland Cortinarius w ah teatiahu tacmu s JV10452 Finland Cortinarius balteatialutaceus IK09-751 TYPE Sweden onc rescm ns 3578 TYPE France 1.00 Cortinarius c C. w ah teatoahwu s RH82.98 TYPE France /Phlegmacioides Cortinarius w ah teatoahwu s IK97-761 Sweden roc ataphihu s RH86-90 TYPE France several synonyms, see Table 1 1.00 Cortinarius cp e.g. C. sawuhm toruu Cortinarius cp roc ataphihu s AY669550 Germany 1.00 aesioc oh or IK97-207 Finland 1.00 Cortinarius c Cortinarius caesiocolor IK00-029 TYPE Finland Cortinarius h atoch aric oh or RH1199 TYPE France C. w ae ioh atus IB53/10 TYPE Germany (= C. eu rus s. auct.) Cortinarius w ae ioh atus IK98-1029 Sweden sihl ae IK95-341 Finland 1.00 Cortinarius areni. Cortinarius areni. sihl ae CFP461b TYPE Sweden Cortinarius myrtilliphilus IK97-1469 TYPE Finland Cortinarius cy rtihh iphihu s TEB 4-94/AY669518 Norway Cortinarius w ah teatiwuhw osus DBB33060 Bulgaria Cortinarius w ah teatiwuhw osus H6027358 Finland Cortinarius w ah teatiwuhw osus IK93-639 Finland 1.00 Cortinarius balteatibulbosus IK98-1624 TYPE Finland Cortinarius w ah teatiwuhw osus IK04-044 Finland Cortinarius w ah teatiwuhw osus AT2004091 Sweden 1.00 Cortinarius w ah teatiwuhw osus AT2004045 Sweden Cortinarius w ah teatiwuhw osus AT2004088 Sweden Cortinarius pseue onewuh aris G42 TYPE France Cortinarius lm neris 2952 TYPE France Cortinarius qh alm scm ntipes Cortinarius qh alm scm ntipes 3603 TYPE France (= C. w ah teatocuc atih is C. w ah teatus var. praestantoiem s PML760 TYPE France Cortinarius ch arow ah toiem s var. h onb ispercu s 833 TYPE France runneol ioh acmu s 2951 TYPE France 1.00 Cortinarius w Cortinarius w runneoh il ieu s Cortinarius w runneoh il ieu s 3734 TYPE France Cortinarius w ah teatotocm ntosus RH306 TYPE France Cortinarius balteatus CFP940 TYPE Sweden Cortinarius w ah teatus 0.99 Cortinarius suw opicu s 3477 TYPE France Cortinarius w runneiaurantius IK93-644 Finland 1.00 Cortinarius brunneiaurantius JV17979 TYPE Finland 1.00 Cortinarius w runneiaurantius JV17979 Finland 0.59 Cortinarius sow rius PAK3235 TYPE Finland 0.98 Cortinarius sow rius IK04-045 Finland s. auct.) Fig. 1 (cont) 116 Persoonia – Volume 33, 2014 0.60 0.86 1.00 0.98 0.98 1.00 1.00 0.74 0.67 0.82 0.56 0.97 0.63 0.55 0.65 Cortinarius ac id ophilu s TEB61-79 TYPE Norway Cortinarius m acul atipe s 2845 TYPE France rassoide s 363 TYPE France 0.95 Cortinarius subc /Phlegmacioides Cortinarius pseud onaev osus Cortinarius sphagne toruu 3746 TYPE France Cortinarius v acc iniophilu s TEB17-88 TYPE Norway Cortinarius pseud onaev osus RH162 TYPE France Cortinarius TYPE Sweden 1.00 Cortinarius pini CFP394 TYPE Norway Cortinarius pini IK90-2288 Finland itriole ns IB97/122 TYPE U.S.A. Wyoming 1.00 Cortinarius c Cortinarius c itriole ns AF325607/IB19970154 U.S.A. Wyoming Cortinarius riousse tia RH84.70-78 TYPE France 1.00 Cortinarius argutus s. auct. AY669535 Norway Cortinarius argutus s. auct. UDB000138 Norway C. raudul f osus var . patricke nsis IB95/617 TYPE U.S.A. California C. patrickensis Cortinarius rosargutus IK08-565 Finland Cortinarius rosargutus IK07-242 Finland Cortinarius rosargutus RH70477 TYPE France Cortinarius rosargutus IK96-1279 Germany /Arguti Cortinarius f raudul osus UDB001036 Italy 1.00 Cortinarius f raudul osus UDB011906 Estonia Cortinarius IK95-1852 TYPE Germany Cortinarius subf raudul osus CFP481 Sweden Cortinarius subf raudul osus IK98-2245 Sweden 0.83 Cortinarius subf raudul osus IK88-2016 Sweden Cortinarius subf raudul osus UDB011261 Estonia Cortinarius subfraudulosus IK11-006 TYPE Norway Cortinarius reve re nd issimu s 4667 TYPE France Cortinarius varius CFP801 TYPE Sweden 0.76 Cortinarius ruf ior 1118 TYPE France Cortinarius v arius Cortinarius saginoide s 1264 TYPE France /Variosimiles Cortinarius pseud opimu s 4516 TYPE France Cortinarius pseud opansa G4401 TYPE France Cortinarius subv ariif orm is 4663 TYPE France C.te olu ingul c atus AB91-10-260 TYPE France Cortinarius v ariosim il is FJ717596 Canada British Columbia C. ariosim v il is IB89/493 TYPE U.S.A. Washington iebe a TYPE U.S.A. Oregon 1.00 Cortinarius w Cortinarius w iebe ae FJ440879 U.S.A. Oregon Cortinarius c alyp trode rmu s 15356 TYPE U.S.A. Michigan Cortinarius c rue ntipell is IK01-053 Estonia 1.00 Cortinarius cruentipellis TN03-1451 TYPE Sweden Cortinarius c rue ntipell is IK11-008 Norway Cortinarius joseph ii 5193 TYPE France 1.00 Cortinarius grac il ior Cortinarius grac il ior JV19538 Italy 0.66 Cortinarius grac il ior UDB001082 TYPE Germany Cortinarius grise oc oe ruleu s IB95/685 TYPE U.S.A. California 1.00 Cortinarius gratus PML85 TYPE France Cortinarius le onic ol or 1.00 Cortinarius anse rinus CFP852 Belgium (= C. anse rinus s. auct., C. am o n e ole ns s. auct.) Cortinarius le onic ol or 4739 TYPE France az onianus TN04-1106 Italy 1.00 Cortinarius pal Cortinarius pal az onianus FJ946938 Spain 0.79 1.00 Cortinarius d iony sa s. auct. AY174813 Germany 1.00 Cortinarius d iony sa s. auct. DQ083782 Germany 0.70 C. iony dsa var. avell aneu s AB97-10-361 TYPE France 1.00 /Dionysae s. lato C. iony dsa var. avell aneu s IK94-1745 France Cortinarius m ahiqu sii e FM202139 TYPE Spain Cortinarius ol iv ace od iony sa IK11-013 Sweden 0.83 Cortinarius ol iv ace od iony sa TN06-306 Finland /Dionysae 1.00 Cortinarius ol iv ace od iony sa IK12-001 Estonia Cortinarius ol i a v od ce i ony s a IK94-1899 France 0.75 1.00 Cortinarius ol iv ace od iony sa TN06-311 Finland Cortinarius ol iv ace od iony sa IK07-1417 Sweden Cortinarius ol iv ace od iony sa IK11-012 Sweden Cortinarius ol iv ace od iony sa MK2540 Finland Cortinarius ol iv ace od iony sa AY669523 Germany ore id iony sa IK07-245 Finland 1.00 Cortinarius b Cortinarius boreidionysaeIK97-1220 TYPE Finland ol orans 4403 TYPE France Cortinarius subdec ol orans 1.00 Cortinarius subdec (= C.tif o mul rm iuu ) Cortinarius subdec ol orans IK11-011 Norway 1.00 Cortinarius ophiopus UDB000729 Sweden Cortinarius ophiopus TYPE U.S.A. Maryland (= C. triumph ans s. auct.) Cortinarius ophiopus IK01-050 Finland Cortinarius kytoevuorii TN05-158 TYPE Finland 1.00 Cortinarius sub rugul osus AB0510263 TYPE France 0.99 Cortinarius sub rugul osus Cortinarius sub rugul osus IK98-2578 Sweden Cortinarius subful igineu s AB97-10-339 TYPE France Cortinarius sp. IK03-005 Sweden Cortinarius c alyp tratus AHS8352 TYPE U.S.A. California sol itarius AHS15377 TYPE U.S.A. Michigan 1.00 Cortinarius sub Cortinarius gl auc oceph alu s IB95/679 TYPE U.S.A. California 1.00 Cortinarius gl auc oceph alu s HQ604682 Canada British Columbia Cortinarius atroch alyb aeu s IB95/630 TYPE U.S.A. California ol iv asce ns AHS14311 TYPE U.S.A. Washington 1.00 Cortinarius sub Cortinarius sub ol iv asce ns AY356323 U.S.A. Washington am aric atus AB94-10-313 TYPE France 1.00 Cortinarius sub Cortinarius tirol ianus Cortinarius tirol ianus PML4341 TYPE France /Glaucopodes Cortinarius al t ic a udu s PML1632 TYPE France 1.00 Cortinarius al tic audu s IK09-1402 Finland Fig. 1 (cont) K. Liimatainen et al.: Identification and nomenclature of Phlegmacium 0.61 1.00 0.58 0.67 1.00 117 1.00 C. glaucopus var. olivaceus f. ingratus PML762 TYPE France Cortinarius scaurocaninus RH71678 TYPE France Cortinarius scaurocaninus HS031095 Germany (= C. magicus s. auct.) Cortinarius pansa IK97-1914 Finland 1.00 Cortinarius pansa AM084701 United Kingdom Cortinarius pansa DQ179120 Sweden Cortinarius pansa IK90-1826 TYPE Finland Cortinarius subfoetens IK08-2010 Finland Cortinarius subfoetens IB89/307 TYPE U.S.A. Wyoming Cortinarius subfoetens GQ159817 Canada British Columbia Cortinarius glaucopoides MICH10358 TYPE U.S.A. Colorado 1.00 Cortinarius glaucopus FJ039615 Canada British Columbia Cortinarius glaucopus CFP786 TYPE Sweden Cortinarius glaucopus Cortinarius glaucopus IK92-1105a Finland Cortinarius glaucopus FJ039616 Canada British Columbia Cortinarius glaucopus EU645632 Canada British Columbia Cortinarius glaucopus var. subrubrovelatus AB97-11-431 TYPE France C. subrubrovelatus 1.00 Cortinarius subrubrovelatus UDB011262 Estonia Cortinarius subrubrovelatus AY174787 Germany Cortinarius misermontii IK98-2417 Sweden Cortinarius misermontii RH84.134 TYPE France 1.00 Cortinarius misermontii IK87-2172 Spain Cortinarius olidoamarus var. valentinus 452173 TYPE Spain Cortinarius foetens IB51/128 TYPE Austria 1.00 /Caerulescentes 1.00 Cortinarius caerulescens CFP853 TYPE Belgium Cortinarius caerulescens AY174863 Germany Cortinarius cremeiamarescens JN889840 U.S.A. Alaska Cortinarius cremeiamarescens IK00-027 Finland 1.00 Cortinarius cremeiamarescens TN06-273 Finland Cortinarius cremeiamarescens IK11-014 TYPE Sweden Cortinarius cremeiamarescens EF218754 Canada British Columbia 1.00 Cortinarius volvatus AHS8857 TYPE U.S.A. California Cortinarius albescens AHS17522 TYPE U.S.A. Washington Cortinarius gentianeus IK11-015 Sweden Cortinarius gentianeus AY669519 Germany 1.00 Cortinarius gentianeus IK97-1378 Finland Cortinarius gentianeus (= C. caesiostramineus s. auct.) Cortinarius gentianeus 2575 TYPE France Cortinarius gentianeus CFP775 Sweden Cortinarius gentianeus IK09-1525 Finland and the reactions were analysed by ABI 3730 DNA Analyzer (Applied Biosystems) automatic sequencer. Sequences were assembled and edited with Sequencer 4.1 (Gene Codes, Ann Arbor, Michigan, USA). A total of 405 new ITS sequences were produced for this study. Collections and GenBank sequences used for the phylogenetic analysis are given in Table 1 and 2. The alignment of 461 ITS sequences was produced with the MUSCLE program (Edgar 2004) under default settings and followed by manual adjustments in BioEdit (www.mbio.ncsu.edu/ BioEdit/bioedit.html). The alignment is 812 nucleotides long (including gaps) and is available at TreeBASE under S14832 (http://www.treebase.org/treebase-web/home.html). Bayesian inference (BI) was performed with MrBayes v. 3.1.1 (Ronquist & Huelsenbeck 2003). The best substitution model for alignment was estimated by both the Akaike information criterion and the Bayesian information criterion with jModelTest 0.1.1 (Posada 2008). GTR model was chosen. Two independent runs with four chains in each were performed 6 000 000 generations with sampling every 100th generation. All trees sampled before stationarity were discarded with a 25 % safety margin (burn-in of 15 000 trees, 1 500 000 generations). Sampled trees from both runs were combined in a 50 % majority rule consensus phylogram with posterior probabilities (PP). The analyses were run with computer clusters of the CSC, IT Center for Science, Espoo, Finland. Morphological study Morphological descriptions are based on material collected by the authors and D. Bojantchev, T.E. Brandrud, T.S. Jeppesen, E. Sesli and A.F. Taylor including specimens in all stages of development. Microscopic characteristics were observed from dried material mounted in Melzer’s reagent (MLZ). Measurements were made in MLZ with an ocular micrometer using 100× oil-immersion lens. Basidiospores were measured from the veil or top of the stipe, 20 spores from one basidiocarp unless otherwise indicated. The length and width were meas- /Glaucopodes Fig. 1 (cont) ured for each spore, and their length/width ratios (Q value) were calculated. The average of all measurements is marked in italic. The lamellar trama and basidia also were examined and the pileipellis structure was studied from scalps from the pileus centre. RESULTS Phylogenetic analyses The 50 % majority rule phylogram resulting from the BI analysis is shown in Fig. 1. Most species are supported by 1.00 PP (or slightly less). Six species received support below 0.95 PP: C. castaneicolor, C. collocandoides, C. flavescentipes (= C. bal­ teatocumatilis s.auct.), C. herpeticus, C. infractiflavus and C. multiformis. Three species, C. balteatoalutaceus, C. pseudo­ cephalixus and C. subfoetens did not form monophyletic groups in our phylogenetic analysis. Most of these nine species represent species that differ from their closest relatives by less than 10 substitutions and indel positions in ITS regions, but they all, however, have low intraspecific variation. In a majority of the species, the intraspecific variation is from 0 to 1 substitutions or indel positions. In some species it is 2 substitutions and/or indel positions. Similarly, in a majority of the species, the interspecific difference is more than 10 substitutions and indel positions, but in some species it is only four, i.e. in the species pair C. claricolor /C. rex­claricolorum. All the species, however, have a clear barcoding gap between intra- and interspecific variation. In the following species the intraspecific variation was more than two substitutions and/or indel positions, but no clear grouping was obtained in the analysis of ITS sequences or the number of specimens was too low for making conclusions: C. albescens /C. gentianeus / C. volvatus complex, C. aureofulvus s.auct. /C. orichalceus var. xanthocephalus complex, C. herpeticus /C. violaceonitens complex, C. misermontii /C. olidoamarus var. valentinus /C. sub­ 118 accedens /C. van­campiae complex, C. pallidirimosus, C. por­ phyropus and C. scaurus. A total of 236 types representing 154 species were successfully sequenced. Of these, 114 species are described only once whereas 40 species had one or more synonyms. The species with the largest number of synonyms are C. largus (14 synonyms), C. pseudonaevosus (6 synonyms), C. talus (5 synonyms), C. crassus (4 synonyms) and C. varius (4 synonyms). All the names of the types are listed in alphabetical order in Table 1, followed by the current name. Several infrageneric groups with three or more species were supported by 1.00 PP: /Calochroi & Fulvi, / Claricolores, /Cyanites, /Dionysae (s.lat. 0.56 PP), /Glaucopodes, /Infracti, /Multiformes, / Phlegmacium, / Purpurascentes and / Scauri. In addition, clade /Arguti (0.98 PP) received some support and clades / Elastici & Percomes (0.94 PP, s.l. 0.53 PP), / Phlegmacioides (0.77 PP) and / Variosimiles (0.74 PP) low support. Taxonomy Persoonia – Volume 33, 2014 Epitype. Belgium, Brabant, Tervuren, in beech forest on calcareous soil, 23 Sept. 1989, Brandrud et al. CFP853 (S), designated here. MycoBank MBT176395. GenBank KF732271. Illustrations — Brandrud et al. (1992: pl. B51), also see MycoBank. Descriptions of the species — Brandrud et al. (1992: pl. B51), Jeppesen et al. (2012: 793). Cortinarius claricolor (Fr.) Fr., Epicr. Syst. Mycol.: 257. 1838 Basionym. Agaricus multiformis δ claricolor Fr., Observ. Mycol. 2: 65. 1818. = Cortinarius pseudoturmalis Bidaud & Moënne-Locc., Atlas des Cortinaires 19: 1503. 2010. Neotype. Sweden, Ångermanland, Stigsjö, Uland, Långmyrberget, in spruce forest with blueberry, 9 Aug. 1988, Brandrud et al. CFP691 (S), designated here. MycoBank MBT176396. GenBank KF732283. Illustrations — Bidaud et al. (2010: pl. 759), Brandrud et al. (1992: pl. B48), Fries (1867–1884: pl. 141). Descriptions of the species — Brandrud et al. (1992: pl. B48), Jeppesen et al. (2012: 821). Neo­ and epitypifications Neotypes for 15 species described by Fries and six species described by Albertini, Batsch, Britzelmayr, Persoon, Schaeffer and Schweinitz are proposed as well as epitypes for three species described by Batsch, Cordier and Schaeffer in the 19th century. These include names that have been commonly used in Europe during the last 20 years (Brandrud et al. 1990,1992, 1994, 1998, Jeppesen et al. 2012) and where the current use of the names is not in contradiction with the protologue. Citations of illustrations and descriptions of the species are provided. If our observations on the species deviate from those of Brandrud et al. (1990,1992, 1994, 1998) and/or Jeppesen et al. (2012), they are presented in comments under each species. For C. cyanites and C. fraudulosus full descriptions are provided since the current use of the name included several closely related species. Synonyms are based on DNA studies of the type specimens and the information on the types is presented in Table 1. The reasonings for synonymy are presented in the discussion. Cortinarius balteatus (Fr.) Fr., Epicr. Syst. Mycol.: 257. 1838 Basionym. Agaricus balteatus Fr., Observ. Mycol. 2: 138. 1818. = Cortinarius subbalteatus Kühner, Bull. Mens. Soc. Linn. Lyon 24, 2: 40. 1955. = Cortinarius balteatotomentosus Rob. Henry ex Rob. Henry, Bull. Soc. Mycol. France 101, 1: 4. 1985. = Cortinarius subopimus Bidaud, Atlas des Cortinaires 7: 231. 1995. Neotype. Sweden, Ångermanland, Säbrå, Överdal, under Picea on cultivated area, 3 Aug. 1990, Brandrud et al. CFP940 (S), designated here. MycoBank MBT176298. GenBank KF732262. Illustrations — Brandrud et al. (1994: pl. C60), Fries (1867– 1884: pl. 142). Descriptions of the species — Brandrud et al. (1994: pl. C60), Jeppesen et al. (2012: 814). Notes — Cortinarius balteatotomentosus was first described in 1958 without indicating a type, nonetheless Index Fungorum and MycoBank report it as valid. The validation has been performed later by Henry (1985) giving the holotype no. 306. Cortinarius caerulescens (Schaeff.) Fr., Epicr. Syst. Mycol.: 265. 1838 Basionym. Agaricus caerulescens Schaeff., Fung. Bavar. Palat. 4: 17. 1774. Lectotype. J.C. Schaeffer, Fung. Bav. I, t. 34, f. I, II, III (1762) (designated in Cortin. Fl. Photogr. II (Swedish version): 11, 1992). Cortinarius cumatilis Fr., Epicr. Syst. Mycol.: 269. 1838 Neotype. Sweden, Närke, Hidinge, Garphyttans National Park, N of the road, grass-herb forest with Corylus, Populus tremula, Ulmus, Fagus, Quercus and Picea, 150 m asl, 20 Sept. 1998, I. Kytövuori 98-2164 (H; NY isoneotype), designated here. MycoBank MBT176397. GenBank KF732293. Illustrations — Brandrud et al. (1990: pl. A47), Fries (1867– 1884: pl. 146). Descriptions of the species — Brandrud et al. (1990: pl. A47), Jeppesen et al. (2012: 824). Cortinarius cyanites Fr., Epicr. Syst. Mycol.: 279. 1838 = Cortinarius pseudocyanites var. paucus Reumaux, Atlas des Cortinaires 15: 1032. 2005. = Cortinarius subcyanites Bidaud, Atlas des Cortinaires 15: 1032. 2005. Neotype. Sweden, Uppland, Uppsala, Stadsskogen, mixed forest, 26 Aug. 2005, A. Taylor 2005069 (S; UPS isoneotype), designated here. MycoBank MBT176398. UNITE No. UDB002193, GenBank KF732355. Illustrations — Bidaud et al. (2005: pl. 534), Fries (1867–1884: pl. 152). Pileus 3–9 cm broad, convex with persistently incurved margin, plano-convex when old, innately fibrillose, greyish blue to greyish brown at centre, greyish blue towards the margin, first viscid but soon dry. Lamellae emarginate, medium spaced to almost crowded, first dark violet, later brownish violet. Stipe 4.5–10 cm long, 1–1.5 cm thick at apex, 1.5–4 cm at base, clavate, blue to greyish blue. Universal veil greyish to brownish grey, forming complete and incomplete fibrillose girdles on stipe. Context bluish white in pileus, stronger blue adjacent to lamellae, bluish in stipe, becoming vinaceous red on exposure, marbled hygrophanous. Odour pleasant, fruity (according to Bidaud et al. 2005). Exsiccata: pileus pale bluish grey to lilac grey, when young with the same tint as C. traganus, often with a pale brownish tint at the centre, weakly fibrillose, stipe bluish grey, brownish at the base. Spores 8.8 – 9.8 –10.9 × 5.2 – 5.7– 6.3 µm, av. = 9.5 –10.0 × 5.5–5.8 µm, Q = 1.59–1.72–1.84, Qav. = 1.66–1.77 (6 specimens, 240 spores), amygdaloid, with a fairly narrow apex, moderately verrucose, some with few small golden yellow guttules, fairly faintly dextrinoid. Basidia 32–41 × 7–9 µm (80 basidia), 4-spored, narrowly clavate, with blood red guttules, many with yellowish contents. Lamellar trama hyphae yellowish, smooth, with abundant small to large to worm-like blood red guttules. Stipe apex hyphae almost colourless to yellowish, smooth, with abundant small to large to worm-like, blood red guttules, the K. Liimatainen et al.: Identification and nomenclature of Phlegmacium guttulate layer thick. On the surface few bands of entangled, ochraceous hyphae mostly without guttules. Pileipellis: epicutis very weakly gelatinous, uppermost hyphae 3–10 µm wide, very pale ochraceous brownish, mostly very sparsely, spot-like incrusted, often with scanty to abundant small blood red guttules, lower down 3–9 µm wide, almost colourless, smooth hyphae with abundant small to large to worm-like, blood red guttules. Hypoderm not developed. Ecology & Distribution — In mixed forests of coniferous and deciduous trees, host unknown. Notes — We studied the C. cyanites species complex and recognized three different species, C. cyanites s.str., C. bo­ reicyanites and C. violaceorubens. All the species were well supported in our phylogenetic analysis and differ from one another by more than 20 substitutions and indel positions. The most distinct of the species is C. violaceorubens. It has a dark, violaceous-brownish pileus, the exsiccata are dirty violaceous grey to violaceous brown, and the spores are largest of the group (av. 9.9–11.0 × 6.3–6.6 µm). It grows in Picea dominated forests, often on rich soil and is known from France, Germany, Sweden and Finland. Sister species C. cyanites and C. boreicyanites both have smaller spores and paler exsiccata. Of these, C. boreicyanites is so far only known from the middle boreal zone of Sweden, Finland and Scotland while C. cyanites has a more southern distribution extending from South Finland and middle Sweden to France. In the protologue of C. cyanites Fries described a species with a “pileo pallide coeruleo”. In addition, the picture of C. cyanites in Fries (1867–1884: pl. 152) illustrates a species with a bluish grey pileus. Attached at the base of the stipe there are leaves of Quercus and Betula and spruce needles. Based on the protologue, illustration, and ecology and distribution of the three species we conclude that the species best fitting the description of Fries is the one described here as C. cyanites and we here propose collection A. Taylor 2005069 as neotype for the species. Cortinarius fraudulosus Britzelm., Ber. Naturhist. Vereins Augsburg 4: 122. 1885 Neotype. Germany, Baden-Württemberg, Freudenstadt, Heiligenbronn, gently sloping grass-herb conifer forest on calcareous soil, Abies alba, Picea abies, Fagus sylvatica, 5 Oct. 1995, I. Kytövuori 95-1852, H7019563 (H; NY isoneotype), designated here. MycoBank MBT176399. GenBank KF732518. Illustration — Abarenkov et al. (2010: photo under the accession no. UDB011906). Pileus 4–8 cm broad, hemispherical to convex, then planoconvex, fibrillose, white to ochraceous white when young, with age becoming pale brownish. Lamellae emarginate, almost distant, first white to very pale brownish grey, later pale brown. Stipe 5–10 cm long, 1–2 cm thick at apex, 1.5–3.5 cm wide at base, clavate to almost bulbose, sometimes slightly rooting, whitish, with handling and with age becomes brownish. Uni­ versal veil white, forming distinct girdles on stipe, sometimes floccose. Context white. Odour not recorded. Exsiccata pale greyish brown. Spores 12.9–13.7–15.0 × 7.3–8.1–8.8 µm, Q = 1.60–1.70 – 1.78 (1 specimen, 20 spores), amygdaloid, with a narrow apex, fairly strongly verrucose, warts anastomosing, not high, some with dark intracellular granules, moderately dextrinoid. Basidia 42–53 × 10–12 µm (20 basidia), 4-spored, clavate, otherwise colourless but with a few dark granular bodies. Lamellar trama hyphae sand brown, full of small dark granules or particles. Stipe apex hyphae yellow brown to red brown, entangled, otherwise colourless, but with small to large to long bodies of red brown to red blackish to black granules, uppermost hyphae 5 –10 µm wide. Pileipellis: epicutis fairly weakly gelatinous, 119 hyphae 5–15 µm wide, ochraceous brownish, finely, densely incrusted, with scanty red brown granules. Hypoderm present. In the overall view red brown and unstructured when seen from above. Ecology & Distribution — In montane and hemiboreal conifer forests, calcicolous. Known from Germany, Italy and Estonia. Fruiting in autumn. Notes — The material of C. fraudulosus formed two well supported groups in our phylogenetic analysis. One group consisted of specimens collected from Germany, Italy and Estonia while the other group included mainly specimens from northern Europe. Since C. fraudulosus Britzelm. is described from Germany, the neotype is chosen among the Central/ southern European clade and the other is described below as a new species C. subfraudulosus. Cortinarius glaucopus (Schaeff.: Fr.) Gray, Nat. Arr. Brit. Pl. 1: 629. 1821 Basionym. Agaricus glaucopus Schaeff., Fung. Bavar. Palat. 4: 23. 1774: sanctioned in Fr., Syst. Mycol. 1: 224. 1821. = Cortinarius glaucopoides Kauffman, Papers from the Michigan Academy of Science, Arts and Letters 1: 133. 1923. Neotype. Sweden, Medelpad, Alnö, Ås brygga, in dry spruce forest on calcareous soil, 21 Sept. 1988, Brandrud et al. CFP786 (S), designated here. MycoBank MBT176400. GenBank KF732315. Illustration — Brandrud et al. (1994: pl. C30). Description of the species — Jeppesen et al. (2012: 802). Notes — The description of Agaricus glaucopus in Fries (1821) is short but fits our species. In addition, an unpublished plate of C. glaucopus painted with the supervision of Fries (S, 0318) exists. It represents a species with red brown pileus and bluish lamellae. It is very similar to the photograph of collection CFP786 in Brandrud et al. (1994), which we propose as a neotype for the species. Our observations of C. glaucopus are consistent with those of Jeppesen et al. (2012), only the length of the spores is slightly different: our measurements 7.3–8.1–9 × 4.5–5.0–5.5 µm, Q = 1.54–1.63–1.76, Jeppesen et al. (2012): 7–8.5 × 4.5–5.5 µm. Cortinarius herpeticus Fr., Epicr. Syst. Mycol.: 268. 1838 = Cortinarius thalliopurpurascens Rob. Henry, Doc. Mycol. 25 (no. 97): 48. 1995. Neotype. Sweden, Ångermanland, Säbrå, Hällenyland, under Picea on cultivated area, 20 July 1990, Brandrud et al. CFP936 (S), designated here. MycoBank MBT176401. GenBank KF732321. Illustration — Brandrud et al. (1994: pl. C08, as C. scaurus var. herpeticus). Descriptions of the species — Brandrud et al. (1994: pl. C08), Jeppesen et al. (2012: 784). Notes — An unpublished plate of C. herpeticus (S, 0324) painted with the supervision of Fries exists. The basidiomes in the illustration are similar to the ones in the photo of collection CFP936 (Brandrud et al. 1994), which we propose as a neotype for the species. Cortinarius herpeticus is distinguished from C. scaurus by a stouter appearance, paler colours, more strongly ornamented spores and above all the lack of sepia-coloured pigments in the epicutis. Cortinarius violaceonitens is fairly similar, but is separated by the spores, which are narrower (9.3–10.0 –10.7 × 5.4– 5.9–6.3 µm), amygdaloid-fusoid, with a shallow suprahilar depression, a low ventral humb and blunt apex, and very strongly verrucose surface especially at the apex. The spores of C. herpeticus are 9.1–10.3 –11.6 × 5.7– 6.4 –7.0 µm, narrowly oblong-ellipsoid, and moderately to strongly verrucose. 120 Cortinarius infractus (Pers.: Fr.) Fr., Epicr. Syst. Mycol.: 261. 1838 Basionym. Agaricus infractus Pers., Observ. Mycol. 2: 42. 1800 (1799): sanctioned in Fr., Syst. Mycol. 1: 223. 1821. = Cortinarius amarocaerulescens Bidaud, Atlas des Cortinaires 18: 1376. 2009. = Cortinarius infractus var. aeruginosus Reumaux, Atlas des Cortinaires 18: 1376. 2009. Neotype. Sweden, Bohuslän, Tossene, Anneröd, beech forest, medium rich soil, 15 Sept. 1986, Brandrud et al. CFP495 (S), designated here. MycoBank MBT176402. GenBank KF732325. Illustrations — Bidaud et al. (2009: pl. 739), Brandrud et al. (1990: pl. A09). Description of the species — Brandrud et al. (1990: pl. A09). Cortinarius largus Fr., Epicr. Syst. Mycol.: 259. 1838 = Cortinarius cephalixolargus Rob. Henry, Bull. Trimestriel Soc. Mycol. France 93, 3: 323. 1977. = Cortinarius clarus Reumaux, Atlas des Cortinaires 8: 291. 1996. = Cortinarius claviceps Reumaux, Atlas des Cortinaires 8: 291. 1996. = Cortinarius congeminus Moënne-Locc. & Reumaux, Atlas des Cortinaires 7: 228. 1995. = Cortinarius cupreoviolaceus Bidaud & Reumaux, Atlas des Cortinaires 8: 292. 1996. = Cortinarius largusiellus Reumaux, Atlas des Cortinaires 8: 293. 1996. = Cortinarius lilacinicolor Reumaux, Atlas des Cortinaires 8: 294. 1996. = Cortinarius lintrisporus Reumaux, Doc. Mycol. 27, no. 106: 53. 1997. = Cortinarius lividoviolaceus Rob. Henry, Doc. Mycol. 17, no. 68: 27. 1987. = Cortinarius occultus Moënne-Locc. & Reumaux, Atlas des Cortinaires 8: 295. 1996. = Cortinarius patibilis var. scoticus Brandrud, Edinburgh J. Bot. 54, 1: 114. 1997. = Cortinarius paracrassus Reumaux, Atlas des Cortinaires 7: 230. 1995. = Cortinarius paracyanopus Moënne-Locc. & Reumaux, Atlas des Cortinaires 8: 296. 1996. = Cortinarius subspadiceus Reumaux, Atlas des Cortinaires 8: 298. 1996. Neotype. Finland, Varsinais-Suomi, Turku, Ruissalo, deciduous forest of Quercus robur, Corylus avellana and some Betula on mull soil, 3 Sept. 2008, K. Liimatainen & T. Niskanen 08-060, H6001957 (H; NY isoneotype), designated here. MycoBank MBT176403. GenBank AB859985. Illustration — Brandrud et al. (1998: pl. D22). Descriptions of the species — Brandrud (1998), Brandrud et al. (1998: pl. D22), Jeppesen et al. (2012: 815). Notes — Based on morphological and molecular data C. largus seems like a uniform species. The collection CFP1085 (Brandrud et al. 1998), however, is from France and therefore not ideal as a type for a species described from Sweden. We do not have our own, well-documented specimen from Sweden and therefore we propose the collection K. Liimatainen & T. Nis­ kanen 08-060 from hemiboreal deciduous forest from south western Finland as a neotype for the species. Cortinarius multiformis Fr., Epicr. Syst. Mycol.: 263. 1838 Neotype. Sweden, Ångermanland, Häggdånger, Sjö, spruce forest with blueberry, 21 Aug. 1986, Brandrud et al. CFP445 (S), designated here. MycoBank MBT176404. GenBank KF732350. Illustration — Brandrud et al. (1990: pl. A45). Descriptions of the species — Brandrud et al. (1990: pl. A45), Jeppesen et al. (2012: 808). Notes — The species as neotypified here fits best the original description of the species and the unpublished plate of C. multiformis (S0350). The reminiscent sister species C. tali­ multiformis, which has been mixed with C. multiformis, has white fibrils on the pileus, and less dextrinoid spores. Our observations of C. multiformis are in concordance with those of Brandrud et al. (1990) and Jeppesen et al. (2012: 808), except that our spore measurements are somewhat larger, 8.6–9.6–10.4 × Persoonia – Volume 33, 2014 5.2– 5.7–6.1 µm, Q = 1.58–1.70 –1.79 than those of Brandrud et al. (1990) and Jeppesen et al. (2012) 8 –9.5 × 5–5.5 µm. Cortinarius pansa (Fr.) Sacc., Syll. Fung. 5: 901. 1887 Basionym. Agaricus pansa Fr., Observ. Mycol. (Havniae) 2: 67. 1818. Neotype. Finland, Varsinais-Suomi, Kemiö, Pederså, at small, abandoned limestone quarries, spruce heath forest, roadside, 35 m asl, 21 Sept. 1990, I. Kytövuori 90-1826 (H; isoneotype NY), designated here. MycoBank MBT176405. GenBank KF732522. Illustration — Fries (1867–1884: pl. 145). Description of the species — Jeppesen et al. (2012: 803). Notes — The description and illustration of C. pansa published by Fries (1818, 1867–1884) fit well with the species presented in Jeppesen et al. (2012). The spore measurements given in Jeppesen et al. (2012) (6–)6.5–7.5 × 4–5 µm differ somewhat from ours 6.8–7.5 –8.2 × 4.3–4.6 –5.0 µm, Q = 1.52–1.64 –1.82. The species was previously included in C. glaucopus but differs from it by more red brown pileus, smaller, less verrucose spores, and habitat often on roadsides, yards, parks and plantations. Cortinarius percomis Fr., Epicr. Syst. Mycol.: 260. 1838 Neotype. Finland, Varsinais-Suomi, Karjaa, Kohagen, herb-rich Picea abies forest with some Corylus avellana, Quercus robur, Betula and Populus tremula, 2 Sept. 2008, K. Liimatainen & T. Niskanen 08-041 (H; isoneotype NY), designated here. MycoBank MBT176406. GenBank KF732380. Illustrations — Brandrud et al. (1994: pl. C56), Fries (1867– 1884: pl. 143). Descriptions of the species — Brandrud et al. (1994: pl. C56), Jeppesen et al. (2012: 812). Notes — Based on morphological and molecular data C. per­ comis seems like a uniform species. The collection CFP1104 (Brandrud et al. 1994), however, is from France and therefore not ideal as a type for a species described from Sweden. We do not have our own, well-documented specimen from Sweden and therefore we propose the collection K. Liimatainen & T. Nis­ kanen 08-041 from hemiboreal Picea abies dominated forest from south western Finland as a neotype for the species. An identical ITS sequence of the species from a specimen collected from Sweden, however, exists in UNITE (UDB000726). Cortinarius porphyropus (Alb. & Schwein.) Fr., Epicr. Syst. Mycol.: 271. 1838 Basionym. Agaricus porphyropus Alb. & Schwein., Consp. Fungorum Lusat.: 153. 1805. = Cortinarius porphyropus var. porphyrophorus Reumaux, Atlas des Cortinaires 18: 1378. 2009. Neotype. Sweden, Jämtland, Ragunda sn, Ragunda, in birch forest on rich soil, 20 Aug. 1988, Brandrud et al. CFP717 (S), designated here. MycoBank MBT176407. GenBank KF732387. Illustration — Brandrud et al. (1992: pl. B55). Descriptions of the species — Brandrud et al. (1992: pl. B55), Jeppesen et al. (2012: 819). Notes — Cortinarius porphyropus is described from Germany. Based on our studies it is a uniform and widespread species occurring at least in Europe and North America (Fig. 1). The most representative collection CFP717 from Sweden is here proposed as a neotype for the species. Cortinarius praestans (Cordier) Gillet, Hyménomycètes: 475. 1874 Basionym. Agaricus praestans Cordier, Champ. France, Discom.: 98. 1870. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium Holotype. Pl. XXI (Cordier, Champ. France, 1870). Epitype. France, Ain, Oyonnax SE, Commune d’Echallon, by the road from St-Germain-de-Joux to Echallon, N of the crossing to Plagne, E sloping, rich Fagus forest with Picea abies, 540 m asl, 27 Oct. 1994, P. & I. Kytövuori 94-1861 (H; isoepitype NY). MycoBank MBT176411. GenBank KF732389. Illustration — Brandrud et al. (1990: pl. A42). Descriptions of the species — Brandrud et al. (1990: pl. A42), Jeppesen et al. (2012: 823). Notes — Cortinarius praestans is one of the most distinctive Cortinarius species and is easy to recognize by its large basidiomata and spores, and habitat with thermophilous deciduous trees. It is described from France, but identical sequences of the species exist from Estonia, Germany, Italy and Sweden (Fig. 1, Table 2). Cortinarius purpurascens Fr., Epicr. Syst. Mycol.: 265. 1838 = Cortinarius eumarginatus Rob. Henry ex Bidaud, Carteret & Reumaux, Atlas des Cortinaires 18, 1, 2: 1378. 2009. Neotype. Sweden, Närke, Hidinge, Garphyttans National Park, S of the road, fairly rich spruce grass-herb forest with Corylus, Populus tremula, Betula and Quercus, 150 m asl, 20 Sept. 1998, I. Kytövuori 98-2121 (H; isoneotype NY), designated here. MycoBank MBT176419. GenBank KF732406. Illustration — Bidaud et al. (2009: pl. 743). Description of the species — Jeppesen et al. (2012: 802). Notes — Based on the data we have so far, the species is known from northern to southern Europe (Fig. 1). Cortinarius russus Fr., Epicr. Syst. Mycol.: 261. 1838 Neotype. Sweden, Ångermanland, Säbrä, Överdal, in dry spruce forest on rich soil, 3 Aug. 1990, Brandrud et al. CFP941 (S), designated here. MycoBank MBT176412. GenBank KF732416. Illustration — Brandrud et al. (1994: pl. C35, C44). Descriptions of the species — Brandrud et al. (1994: pl. C35), Jeppesen et al. (2012: 817). Cortinarius scaurus (Fr.: Fr.) Fr., Epicr. Syst. Mycol.: 268. 1838 Basionym. Agaricus scaurus Fr., Observ. Mycol. 2: 75. 1818. = Cortinarius parolivascens Moënne-Locc. & Reumaux, Atlas des Cortinaires 18: 1375. 2009. = Cortinarius scaurus var. notandus Bidaud, Atlas des Cortinaires 18: 1375. 2009. = Cortinarius scaurus f. phaeophyllus M.M. Moser, Fungi non Delineati 15: 18. 2001. Neotype. Switzerland, Bern, Fribourg, Rechthalten, on the border of a peatbog with Pinus strobus, 29 Sept. 1991, Brandrud et al. CFP1074 (S), designated here. MycoBank MBT176413. GenBank KF732423. Illustrations — Brandrud et al. (1994: pl. C21), Fries (1867– 1884: pl. 146). Descriptions of the species — Brandrud et al. (1994: pl. C21), Jeppesen et al. (2012: 783). Notes — Cortinarius scaurus is a widespread species and to date known from eastern North America and Europe (Fig. 1). The most representative collection CFP1074 from Switzerland is proposed as a neotype and the ITS sequence of the specimen is identical to the UNITE sequence UDB001068 from Femsjö, Sweden. Cortinarius serarius Fr., Epicr. Syst. Mycol.: 269. 1838 Neotype. Sweden, Ångermanland, Häggdånger, Torrom, in dry spruce forest on rich soil, 11 Aug. 1990, Brandrud et al. CFP959 (S), designated here. MycoBank MBT176414. GenBank KF732428. Illustration — Brandrud et al. (1994: pl. C25). 121 Descriptions of the species — Brandrud et al. (1994: pl. C25), Jeppesen et al. (2012: 824). Cortinarius subpurpurascens (Batsch) Fr., Epicr. Syst. Mycol.: 265. 1838 Basionym. Agaricus subpurpurascens Batsch, Elench. Fung., cont. prim.: 71. 1786. = Cortinarius largoides Rob. Henry ex Bidaud, Carteret & Reumaux, Atlas des Cortinaires 18: 1378. 2009. = Cortinarius subinops Reumaux, Atlas des Cortinaires 18: 1379. 2009. Holotype. Batsch, Elench. Fung., cont. prim. tab. 16: 74. 1786. Epitype. Finland, Varsinais-Suomi, Turku, Ruissalo, deciduous forest of Quercus robur, Corylus avellana and some Betula on mull soil, 3 Sept. 2008, K. Liimatainen & T. Niskanen 08-059 (H; isoepitype NY), designated here. MycoBank MBT176420. GenBank KF732449. Illustrations — Bidaud et al. (2009: pl. 749, 750). Description of the species — Jeppesen et al. (2012: 819). Notes — Based on the data we have so far the species is known from northern to southern Europe (Fig. 1). A representative collection Liimatainen & Niskanen 08-059 from South Finland is here proposed as an epitype of the species. Cortinarius subtortus (Pers.) Fr., Epicr. Syst. Mycol. (Upsaliae): 273. 1838 Basionym. Agaricus subtortus Pers., Syn. Meth. Fung. (Göttingen) 2: 284. 1801, sanctioned in Fr., Syst. Mycol. 1: 222. 1821. Neotype. Sweden, Inre Småland, Femsjö, Prästskogen, partly paludified spruce forest with some deciduous tree species, 175 m asl, 18 Sept. 1987, I. Kytö­ vuori 87-1510 (H; isoneotype NY), designated here. MycoBank MBT176415. GenBank KF732454. Description of the species — Jeppesen et al. (2012: 811). Cortinarius talus Fr., Epicr. Syst. Mycol.: 263. 1838 = Cortinarius aurantionapus Bidaud & Reumaux, Atlas des Cortinaires 16: 1096. 2006. = Cortinarius crenulatus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires 16: 1097. 2006. = Cortinarius ochropudorinus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires 16: 1097. 2006. = Cortinarius pseudominor Rob. Henry ex Reumaux, Atlas des Cortinaires 16: 1098. 2006. = Cortinarius pseudotalus Rob. Henry ex Bidaud & Reumaux, Atlas des Cortinaires 16: 1098. 2006. Neotype. Sweden, Jämtland, Ragunda sn, Ragunda, in birch forest on rich soil (Betula, Populus, Pinus), 26 Aug. 1989, Brandrud et al. CFP832 (S), designated here. MycoBank MBT176416. GenBank KF732457. Illustrations — Brandrud et al. (1992: pl. B47), Fries (1867– 1884: pl. 145). Descriptions of the species — Brandrud et al. (1992: pl. B47), Jeppesen et al. (2012: 811). Notes — Our observations of the species are in concordance with those of Brandrud et al. (1992) and Jeppesen et al. (2012), except for the length of the spores, which according to our measurements is 7.3– 8.0 –8.8 × 4.5– 5.0–5.2 µm and in Brandrud et al. (1992) and Jeppesen et al. (2012) 7.5–9.5 × 4.5–5.5 µm. Cortinarius turmalis Fr., Epicr. Syst. Mycol.: 257. 1838 Neotype. Sweden, Medelpad, Borgsjö, Julåsen, in herbaceous spruce forest, 20 Aug. 1988, Brandrud et al. CFP716 (S), designated here. MycoBank MBT176417. GenBank KF732464. Illustration — Brandrud et al. (1994: pl. C31). Descriptions of the species — Brandrud et al. (1994: pl. C31), Jeppesen et al. (2012: 821). 122 Cortinarius variecolor (Pers.: Fr.) Fr., Epicr. Syst. Mycol.: 259. 1838 Basionym. Agaricus variecolor Pers., Syn. Meth. Fung. 2: 280. 1801. Sanctioned in Fr., Syst. Mycol. 1: 222. 1821. = Cortinarius muricinicolor Moënne-Locc., Atlas des Cortinaires 8: 295. 1996. = Cortinarius piriodolens Moënne-Locc., Atlas des Cortinaires 8: 296. 1996. Neotype. Sweden, Gotland, Viklau, Tjaukle, in spruce forest on calcareous soil, 29 Sept. 1990, Brandrud et al. CFP1021 (S), designated here. MycoBank MBT176418. GenBank KF732466. Illustrations — Brandrud et al. (1992: pl. B20), Fries (1867– 1884: pl. 144). Descriptions of the species — Brandrud (1998), Brandrud et al. (1992: pl. B20), Jeppesen et al. (2012: 815). Persoonia – Volume 33, 2014 NEw SpECIES AND COMbINATIONS Species with an isolated position Cortinarius cremeiamarescens Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805786; Fig. 2a, 3a Etymology. The name refers to the colour of the basidiomata and bitter taste of pileus cuticle. = Cortinarius caesiostramineus Rob. Henry sensu Brandrud et al. 1990, Jeppesen et al. 2012, p.p. Type. Sweden, Gotland, Alskog and När parish, Ollajvs Nature Reserve, mesic to damp spruce forest with some Pinus, Quercus and Corylus, 27 Sept. 2011, I. Kytövuori 11-014 (holotype H; isotype NY). GenBank KF732493. Pileus 3.5–5.5 cm broad, hemispherical to convex, then expanded, very finely innately fibrillose, cream-coloured to pale Fig. 2 Photo of: a. C. cremeiamarescens TN06-273; b. C. flavivelatus IK98-885; c. C. kytoevuorii TN05-158; d. C. flavipallens T. Kekki 368; e. C. boreidionysae IK97-1220; f. C. cruentipellis IK01-053; g. C. caesiophylloides TEB277-09. — Photos: a, c. Kare Liimatainen; b, e, f. I. Kytövuori; d. T. Kekki; g. T.E. Brandrud. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium 123 ochraceous yellow. Lamellae emarginate, almost crowded, pale greyish brown. Stipe 6.5–9.5 cm long, 0.7–1.4 cm thick at apex, 1–2.3 cm wide at base, with slightly marginate bulb, at first white, becoming pale brownish yellow with age. Universal veil white, sparse, at bulb margin. Context white. Odour indistinct. Taste: pileus cuticle bitter. Exsiccata: pileus ochraceous clay-colour to ochraceous yellow to warm ochraceous brown especially at the centre, stipe pale grey to brown. kanen et al. 03-1255, H7018363 (H); F03-1163, H7018395 (H); Närke. Hidinge, Garphyttans Nationalpark, herb-rich Picea abies forest with Cory­ lus, Populus tremula, Betula and Quercus, 26 Sept. 2004, T. Niskanen et al. 03-1255, H7017775 (H). In MLZ: Spores 7.0–7.8 –8.8 × 4.3– 4.7–5.0 µm, av. = 7.5–8.3 × 4.6–4.9 µm, Q = 1.54–1.67–1.87, Qav. = 1.59–1.74 (9 specimens, 260 spores, Fig. 3a), citriform to narrowly fusoid, beaked, thin-walled, fairly finely, densely, and often sharply verrucose, slightly to moderately dextrinoid. Basidia 24–34 × 6.5–8 µm (80 basidia), 4-spored, narrowly clavate, very thin-walled, colourless, more or less filled with blood red drops. Lamellar trama hyphae yellow, filled with small blood red drops, larger globose and worm-like guttules fairly scanty (especially more scanty than in C. gentianeus). Stipe apex hyphae almost colourless to yellow, smooth, full of small golden yellow to blood red drops, larger globose and worm-like guttules fewer. Pileipellis: epicutis strongly gelatinous, hyphae 2–6 µm wide, very thin-walled and difficult to define, filled with small to medium-sized blood red guttules, mostly evenly distributed in the hyphae. Hypoderm present, pale yellow. In C. gentianeus the epicutis hyphae are full of very long, worm-like, foamy guttules and the blood red layer is much thicker than in C. cremeiamarescens. Ecology & Distribution — In hemiboreal and southern boreal conifer-dominated forests on rich to calcareous soil. Known from southern Europe and western North America, British Columbia and Alaska. Fruits from late August to late October. Notes — Cortinarius cremeiamarescens was previously confused taxonomically with C. gentianeus Rob. Henry (= C. cae­ siostramineus sensu Jeppesen et al. 2012 p.p.). However, the latter species is larger, typically has paler, more whitish or greyish exsiccata and larger (7.7– 8.5–9.3 × 4.8–5.2 –5.4 µm, Q = 1.54–1.65–1.76), amygdaloid to citriform, less thin-walled, more dextrinoid, and more verrucose spores, and much more abundant large, blood red, foamy, worm-like guttules in the pileipellis and lamellar trama. Cortinarius cremeiamarescens formed a well-supported clade in our phylogenetic analysis (1.00 PP). The ITS sequences of the species are identical and it differs from its sister species C. gentianeus by 17 substitutions and indel positions. Further relationships with other species of Cortinarius were not resolved in our analysis. Other specimens examined. Finland, Varsinais-Suomi, Parainen, Lemlahdensaari, Fallskogen, Pinus sylvestris heath forest with some Picea abies on sandy soil, by the calcareous dust road, 20 Oct. 2006, K. Liimatainen & T. Niskanen 06-273; Uusimaa, Espoo, Luukkaa outdoor recreation area, N of Haukkalampi, mesic, partly grass-herb-spruce forest with some Populus tremula, Betula and Pinus sylvetris, 9 Sept. 2004, K. Liimatainen & T. Nis­ kanen 04-768, H6029461; Kirkkonummi, Dorgarn, Meiko-Trehörningen nature reserve, semi-open spruce forest with some hardwood bushes, 27 Sept. 2007, I. Kytövuori 07-1722, H6001575 (H); Siuntio, Lappträsk, grass-herb-spruce forest with some Pinus, Betula, Populus and Corylus, 24 Aug. 2000, I. Kytövuori 00-027 (H); Etelä-Savo, Mäntyharju, Juolasvesi, Hietaniemi, Sojonkangas, fairly rich spruce-pine forest with abundant Betula and Populus tremula, 29 Sept. 1994, I. Kytövuori 94-1177b, H6035734 (H). – Sweden, Västergötland, Undenäs, c. 1.5 km NNW of Sätra bruk, herb-rich Picea abies forest with some Betula, Populus and Pinus, 6 Sept. 2003, T. Nis­ a Additional specimens. Canada, British Columbia, Interior Cedar Hemlock Forest, mycorrhizal root tip of Betula papyrifera, isolate UBCOCS153F, GenBank EF218754. – USA, Alaska, Delta Junction, source: boreal forest, soil 0–20 cm, GenBank JN889840. Cortinarius flavivelatus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805863; Fig. 2b, 3b Etymology. The name refers to the yellow universal veil. Type. Sweden, Norrbotten, Pajala, Junosuando, Nature Reserve Area between Sarvikero and Tulemajoki, dryish Picea abies heath forest with Pinus, Betula, and with open meadows, 15 Aug. 1998, I. Kytövuori 98-885 (holotype H; isotype NY). GenBank KF732528. Pileus 5–8 cm broad, hemispherical to convex, then plano-convex, viscid, finely innately fibrillose, olive brown to ochraceous brown to brown at the centre, lighter at the margin, with hygrophanous streaks. Lamellae emarginate, crowded, first distinctly pale bluish, later pale brown with a bluish tint. Stipe 6–10 cm long, 1–1.8 cm thick at apex, 1.5–2.5 cm wide at base, clavate to almost cylindrical, whitish, with a bluish tint at the apex. Universal veil yellow, forming girdles on stipe, somewhat viscid. Context white in pileus and lower part of the stipe, bluish at stipe apex. Odour indistinct. KOH­reaction negative in all parts. Exsiccata: pileus warm yellowish to reddish brown, stipe whitish. b c d e Fig. 3 Spores of: a. C. cremeiamarescens TN04-768; b. C. flavivelatus IK98-885; c. C. kytoevuorii TN05-158; d. C. ochribubalinus IK93-641; e. C. subfraudu­ losus IK11-006, in Melzer’s reagent. — Drawn by I. Kytövuori and T. Niskanen. — Scale bar = 10 µm. 124 In MLZ: Spores 9.1– 9.8 –10.9 × 5.0–5.4 –5.9 µm, Q = 1.67– 1.82–1.98 (1 specimen, 60 spores, Fig. 3b), amygdaloid-fusoid to slightly, narrowly citriform, with a shallow suprahilar depression and somewhat beaked apex, fairly finely, separately verrucose, slightly dextrinoid. Basidia 27–38 × 7.5–8 µm (40 basidia), 4-spored, clavate, pale sand brown, with few dark red brown granules, almost hyaline when mature. Lamellar trama hyphae: with abundant dark granules and chips, sometimes in small mounds. Stipe apex hyphae pale yellowish sand brown with small brown granules, outermost hyphae entangled, narrow, with more or less abundant blackish red granules. Pileipel­ lis: epicutis strongly gelatinous, uppermost hyphae 5–10 µm wide, ochre brown, finely to strongly spirally incrusted, mostly not granulose, lower down dark-granulose. Hypoderm well developed, red brown. The upper part of the hypoderm and the transition hyphae towards the epicutis strongly incrusted and with mounds of blackish brown granules. Evenly distributed small granules present (sometimes absent) between the mounds. Ecology & Distribution — In northern boreal coniferous forests. Known from Sweden, Norrbotten. No sequences of this species exist in public databases. Notes — Based on morphology and molecular data C. flavi­ velatus is a sister species of C. pini Brandrud. Cortinarius pini, however, has white, sometimes ochraceous white universal veil and much larger spores (10.7–11.7–12.9 × 6.3–6.8–7.3 µm). In ITS regions the difference between the species is 17 substitutions and indel positions. Cortinarius kytoevuorii Niskanen & Liimat., sp. nov. — MycoBank MB805865; Fig. 2c, 3c Etymology. The species is named in honour of Ilkka Kytövuori, a mycologist from Finland. Type. Finland, Koillismaa, Kuusamo, Oulanka, Ampumavaara, S slope, old, grass-herb Picea abies forest with some Betula, Pinus sylvestris and Populus tremula, on calcareous soil, 17 Sept. 2005, T. Niskanen & K. Liima­ tainen 05-158, H6029355 (holotype H; isotype NY). GenBank KF732529. Pileus 6–9 cm broad, hemispherical to convex, then expanded, finely innately fibrillose, yellow brown to brown, with hygrophanous streaks. Lamellae emarginate, almost crowded to medium spaced, at first pale brownish grey, becoming more brown with age. Stipe 6–9 cm long, 1.2–1.5 cm thick at apex, 2–2.5 cm wide at base, with fairly narrow, marginate bulb, at first white, becoming pale brownish yellow with age. Context whitish to pale yellow. Odour indistinct. KOH reactions: in pileipellis brown (no reddish tints); in context, mycelium and bulb margin negative. Exsiccata: pileus dull, dark red brown overall, stipe almost concolorous with pileus. In MLZ: Spores: 7.5–8.5–9.5 × 5.0–5.3 –5.4 µm, Q = 1.54– 1.61–1.72 (1 specimen, 60 spores, Fig. 3c), amygdaloidellipsoid, very strongly, separately, ± sharply verrucose (C. por­ phyropus like but more dextrinoid), slightly to moderately dextrinoid. Basidia: 24 – 32 × 7.5 – 9 µm (20 basidia), 4-spored, clavate. Lamellar trama hyphae: pale pellucid yellowish, guttulate, smooth. Stipe apex hyphae colourless to pale strawcoloured, smooth, guttulate. Pileipellis: epicutis in overall view orange, (very) weakly gelatinous, hyphae 5–10 µm wide, finely to strongly spirally incrusted, many of the uppermost ones abundantly with intercellular, unevenly distributed, orange red granules, granule mounds and concretions. Large-celled hypoderm present, yellowish and with scanty small orange granules in the upper part, lower down colourless. Ecology & Distribution — In coniferous forests, on calcareous soil. Fruits in autumn. Notes — Cortinarius kytoevuorii is reminiscent of C. glauco­ pus but is more slender, has a yellow brown to brown pileus, Persoonia – Volume 33, 2014 and lacks bluish tints in basidiomata. It is most easily recognised by the orange red granules and concretions in the uppermost hyphae of the pileipellis when mounted in MLZ. The sister species C. subrugulosus Bidaud & Armada has a more southern distribution. The northernmost known collection is from Sweden, Öland under Fagus (Kytövuori 98-2578 (H)). Furthermore, the spores are shorter (7.3–7.9 –8.6 × 5.0–5.3 –5.7 µm), relatively broader (Qav. = 1.5), less verrucose, and more strongly dextrinoid, and in the pileipellis small yellow to blood-red guttules are seen in MLZ. In our phylogenetic analysis C. kytoevuorii formed a well-supported clade (1.00 PP) with C. subrugulosus, but further relationships were not solved. The difference in ITS regions between the two sister species is 11 substitutions and indel positions. Cortinarius ochribubalinus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805866; Fig. 3d Etymology. The name refers to the colour of the basidiomata. Type. Finland, Uusimaa, Espoo, Nuuksio, the open-air territory of Pirttimäki, between the main road and the lake, opposite the parking area, fairly rich grass-herb forest with Populus tremula, Betula, Alnus incana, Quercus, Corylus, Prunus padus, Salix spp., and some old pines and young spruce, 2 Sept. 1993, I. Kytövuori 93-641, H6032734 (holotype H; isotype NY). GenBank KF732530. Pileus 5–8 cm broad, convex, soon plano-convex, sometimes with a broad umbo, very finely fibrillose, centre ochraceous, whitish towards margin. Lamellae emarginate, medium spaced, at first very pale brownish grey, later pale brown. Stipe 6–10 cm long, 0.8–1.3 cm thick at apex, 1.5–2 cm at base, clavate, at first white, becoming pale brownish yellow with age. Universal veil white, on pileus margin thin, forming thin belts on the stipe. Context white. Odour pleasant. Exsiccata: pileus warm yellowish brownish at centre, pale leather-coloured to almost whitish at margin, stipe concolorous with the pileus margin. In MLZ: Spores 12.5–13.4 –14.3 × 7.5–7.9 –8.2 µm, Q = 1.57– 1.69–1.84 (1 specimen, 60 spores, Fig. 3d), amygdaloid, strongly verrucose, most strongly so at the apex, very much like those in C. riederi group or C. violaceus, moderately dextrinoid. Ba­ sidia 39–48 × 9–12 µm (20 basidia), light sand brown, with large to small blood black granules. Lamellar trama hyphae with blood black substance and/or same-coloured granules, dark chips almost lacking. Stipe apex hyphae yellow, with colourless guttules, coloured granules lacking, but outermost hyphae sand brown to somewhat redder, with few to abundant blood red, small granules. Velum/Cortina hyphae sand brown to somewhat redder, with few to abundant blood red, small granules. Pileipel­ lis: epicutis somewhat gelatinous, hyphae 3–10 µm wide, near the surface ochraceous brown, finely, densely incrusted, mostly not granulose, lower down strongly granulose with small to large blackish brown granules in mounds and long, sausage-shaped clusters and concretions. Hypoderm well developed, red brown, hyphae mostly granulose in the upper part. Ecology & Distribution — With deciduous trees, possible hosts Populus, Corylus or Quercus. Known from South Finland. No sequences of this species exist in public databases. Notes — Based on our molecular studies C. ochribubalinus does not have any known, close relatives. It holds an isolated position in the phylogenetic tree and differs in ITS regions by more than 20 substitutions and indel positions from the closest species C. patrickensis (M.M. Moser) Niskanen, Liimat., Kytöv., Bojantchev & Ammirati. Morphologically, it is reminiscent of species in /Arguti. The spores are large and similar to those of C. fraudulosus and C. subfraudulosus, but more strongly verrucose. The smell of the lamellae is also different, pleasant in C. ochrobubalinus and often earthy-raphanoid in C. fraudulosus and C. subfraudulosus. In addition, the universal veil is sparse K. Liimatainen et al.: Identification and nomenclature of Phlegmacium where as C. fraudulosus and C. subfraudulosus have more abundant sometimes even floccose universal veil remnants on stipe. /ARgUTI Cortinarius patrickensis (M.M. Moser) Niskanen, Liimat., Kytöv., Bojantchev & Ammirati, comb. nov. — MycoBank MB805883 Basionym. Cortinarius fraudulosus var. patrickensis M.M. Moser, Mycotaxon 74, 1: 10. 2000. Type. USA, California, Humboldt Co., Patrick’s Point State Park prope Trinidad, in coniferous forest (Picea sitchensis, Pseudotsuga menziesii), 25 Nov. 1995, M. Moser 95/617 (holotype IB). GenBank KF732307. Other specimens examined. Sweden, Medelpad, Borgsjö, Julåsen, SE sloping, partly paludified, rich grass-herb-spruce forest with some pines and birches, 15 Sept. 1995, I. Kytövuori 95-1400, H7019584 (H). – USA, Six Rivers Nat’l forest, off Hwy 299, 1004 m asl, mixed forest (Notholithocarpus densiflorus, Quercus kelloggii, Pinus spp., Pseudotsuga menziesii, etc.), 20 Nov. 2010, D. Bojantchev DBB39406 (UC). Notes — Cortinarius patrickensis is a typical member of /Arguti, it has a whitish to pale brown basidiomata and large spores, and the placement is also supported by molecular data. It was first described as a variety of C. fraudulosus but based on our genetic and morphological data it should be treated as a species. In our phylogenetic analysis C. patrickensis grouped together with C. rosargutus Chevassut & Rob. Henry (0.86 PP). Also with the pairwise comparison the closest species is C. rosargutus from which it differs by 5 substitutions and indel positions in ITS regions. Both taxa, C. patrickensis and C. rosargutus, include several identical or almost identical sequences and have a clear barcoding gap (no overlap between intra- and interspecific variation). A description of the species is provided in Moser & Ammirati (2000). Typical for the species are amygdaloid, large spores 12.2–13.3 –14.7 × 7.0–7.7–8.4 µm, Q = 1.62 –1.73 –1.84, and green apple-like to strong green corn-like odour. The spores are narrower than those of C. ros­ argutus (Qav. = 1.60). Cortinarius patrickensis grows in coniferous forests, often on calcareous soil. It was previously only known from California, USA but is here reported for the first time from Europe in Sweden. Species might be occasional in suitable habitats, at least in Europe and North America, but has most likely been misidentified as either C. fraudulosus or C. rosargutus. Cortinarius subfraudulosus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805867; Fig. 3e Etymology. The name refers to the affinity of C. fraudulosus. Type. Norway, Oppland, Lunner, Skøyenåsen, Picea abies forest with Corylus on calcareous soil, 3 Sept. 2011, I. Kytövuori 11-006 (holotype H; isotype NY). GenBank KF732564. Illustrations — Brandrud et al. (1990: pl. A07 as C. argutus subsp. fraudulosus), Abarenkov et al. (2010: photo under the accession no. UDB011261). Pileus 4–10 cm broad, hemispherical to convex, then planoconvex, finely fibrillose, white to ochraceous white when young, with age becoming pale brownish. Lamellae emarginate, medium spaced to almost distant, first white to very pale brownish grey, later pale brown. Stipe 5–10 cm long, 1–2 cm thick at apex, 1.5–3 cm wide at base, clavate to slightly bulbous, sometimes slightly rooting, whitish, becoming brownish with handling and with age. Universal veil white, forming distinct girdles on stipe, sometimes floccose. Context white, but becomes very slowly black when bruised. Odour weak, a combination of earthy and raphanoid. Exsiccata: pileus dirty greyish to yellow brown 125 at the centre, whitish to yellowish brown towards the margin, stipe greyish white to pale brown. In MLZ: Spores 12.0–13.6–15.0 × 7.0–7.8–8.4 µm, av. = 12.5– 14.4 × 7.6–8.2 µm, Q = 1.57–1.72 –1.88, Qav. = 1.64–1.75 (4 specimens, 100 spores, Fig. 3e), amygdaloid (to weakly citriform), with a narrow apex, moderately to fairly strongly verrucose, warts anastomosing, not high, moderately dextrinoid. Basidia 38–55 × 10–12 µm (2 specimens, 60 basidia), almost colourless to pale sand brownish, clear or with few small, brown granules. Lamellar trama hyphae full of dark granules or chips or blood black particles. Stipe apex hyphae yellow to reddish brown, outermost ones with abundant large red brown to blood blackish granule masses or opaque particles. Velum/ Cortina hyphae with abundant large red brown to blood blackish granule masses or opaque particles. Pileipellis: epicutis fairly weakly gelatinous, hyphae at the surface 3–10 µm wide, finely incrusted, ochraceous brownish, mostly not granulose, lower down somewhat granulose or not. In the transition between epicutis and hypoderm up to 15 µm wide, dark red brown, strongly incrusted hyphae with large, blackish granula mounds, intracellular concretions and/or red brown, 3–15 µm wide extracellular pigment mounds on the hyphae, with few evenly distributed small granules. Hypoderm well developed, red brownish to very dark red brown, somewhat granulose or not. Ecology & Distribution — In hemiboreal and southern boreal coniferous forests, calcicolous. Known from Fennoscandia and Estonia. Fruiting in autumn, in September. Other specimens examined. EStonia, Hiiumaa, Kõrgessaare, Kõpu, old spruce forest with Pinus, Betula, Quercus, Populus tremula and Corylus, on calcareous soil, 15 Sept. 2001, T. Niskanen & I. Kytövuori (H); Pühalepa, Kallaste pank, spruce forest with Pinus, Betula, Populus tremula and Corylus, on calcareous soil, 23 Sept. 2008, J. Vesterholt & J. Vauras 26655F (TURA). – Sweden, Östergötland, Väversunda parish, E sloping, very rich spruce grass-herb forest with hardwood bushes, 26 Sept. 1988, I. Kytövuori 88-2016 (H); Uppland, Börstils sn, Marieberg, Täpporna, in rich spruce forest, 11 Sept. 1986, T.E. Brandrud et al. CFP481, F44801 (S); Södermanland, Mörkö parish, Oaxen, dryish spruce grass-herb forest with Pinus, Betula, Populus tremula and Salix spp., on calcareous soil, 27 Sept. 1998, I. Kytövuori 98-2244 (H), 98-2245 (H). Additional specimen. EStonia, Saare, Kihelkonna, Mäebe, in coniferous forest, 21 Sept. 2009, V. Liiv TU106607, UNITE No. UDB011261 as C. frau­ dulosus (TU(M)). Notes — The species has earlier been called C. fraudulosus in northern Europe. Cortinarius fraudulosus, however, has been described by Britzelmayr (1885) from Siebentischwald, Bavaria, Germany. In our phylogenetic tree the northern material and the more southern European material formed two separate, well-supported clades which differ from one another by 9 substitutions and indel positions in their ITS regions. The sequences of C. subfraudulosus have one base polymorphisms and the maximum pairwise distance is zero. Therefore, we here describe the northern species as new. /CALOCHROI & FULvI Cortinarius flavipallens Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805868; Fig. 2d, 4a Etymology. The name refers to the colour of the pileus. Type. Finland, Kainuu, Kajaani, Hietalahti, Torakangas NNW of Korpitaipale, by the power line, fairly damp grass-herb-spruce forest with Pinus, Betula and Populus tremula, on calcareous soil, 13 Sept. 2008, I. Kytövuori 08-1729, H6032745 (holotype H; isotype NY). GenBank KF732554. Pileus 4–8 cm broad, hemispherical to convex, soon expanded, pale ochraceous to pale brown. Lamellae emarginate, crowded, pale grey to pale greyish brown. Stipe 4–6 cm long, 1–1.5 cm thick at apex, 2–2.5 cm wide at base, with a marginate bulb, white. Universal veil whitish to pale brown at bulb margin. 126 Persoonia – Volume 33, 2014 Context white. Odour indistinct. KOH reactions on pileus and bulb margin red, on basal tomentum and rhizomorphs slowly and/or weakly vinaceous pink. Exsiccata: pileus pale ochre to ochre brown, darkest at the centre, paler towards the margin, stipe greyish white to pale brown. In MLZ: Spores 9.5–10.7–11.6 × 5.4–6.0 –6.3 µm, av. = 10.3– 11.0 × 5.9 – 6.1, Q = 1.65 –1.77–1.85, Qav. = 1.73 –1.80 (3 specimens, 120 spores, Fig. 4a), amygdaloid-fusoid to weakly citriform, with a shallow suprahilar depression and mostly a somewhat blunt apex, moderately verrucose, warts anastomosing or not, slightly to moderately dextrinoid. Basidia 26–38 × 8–10 µm (60 basidia), 4-spored, clavate, very pale yellowish, clear, few slightly granulose-guttulate. Lamellar trama hyphae pale yellowish, smooth, sometimes guttulate, some colourless crystals solitary or in roundish masses, not in branch-like fascicles. Pileipellis: epicutis strongly gelatinous, hyphae on the surface 3–5 µm wide, very pale ochre, smooth to very finely, densely, spirally incrusted, walls distinctly visible, also some hyphae with yellow, foamy contents. Lower down equally wide to slightly wider, strongly incrusted hyphae, often in bundles. Hypoderm absent. In KOH the gelatinized hyphae pale vinaceous pink. Ecology & Distribution — In boreal Picea abies dominated forests on calcareous soil. So far only known from Finland but might be widely distributed, since an ITS sequence (FJ039640) from a specimen collected in British Columbia in western Canada differed only by 4 substitutions and indel positions from Finnish material and might be conspecific. Other specimens examined. Finland, Laatokan Karjala, Parikkala, Vaaranperä, Soininmäki, S part, Soininjoki, fairly old, SE sloping spruce forest with some Pinus, Betula and Populus tremula, 17 Sept. 2009, A. Ahola H6032393 (H); Perä-Pohjanmaa, Tervola, Peura, Raemäki, in Picea dominated forest on calcareous soil, 5 Sept. 2011, T. Kekki 368 (H). Notes — Cortinarius flavipallens with its pale ochraceous colours is in appearance between C. metarius Kauffman (= C. bar­ barorum Bidaud, Moënne-Locc. & Reumaux) and C. caesio­ cinctus Kühner, without the bright yellow pileus of the former or the bluish grey one of the latter. In addition, the KOH reaction in the mycelium is slower and weaker than in the former two species. Cortinarius piceae Frøslev, T.S. Jeppesen & Brandrud and C. corrosus Fr. differ from C. flavipallens by their negative KOH reaction in the basal mycelium. The ITS sequences of C. flavipallens were identical and it formed a well-supported clade in our phylogenetic analysis. However, the relationships with other species of /Calochroi p.p. were not resolved. a d Cortinarius luteicolor (A.H. Sm.) Ammirati, Bojantchev, Niskanen & Liimat., stat. nov. & nom. nov. — MycoBank MB805896 Etymology. The name refers to the yellowish colours of the pileus and lamellae. Basionym. Cortinarius orichalceus var. olympianus f. luteifolius A.H. Sm., Lloydia 7: 185. 1944. Type. USA, Washington, Olympic Mts, near Lake Angeles, 19 Sept. 1941, A.H. Smith 16970, barcode 10389 (holotype MICH). GenBank KF732368. Other specimens examined. USA, California, Sierra Nevada, Yosemite Nat’l Park, Hwy 120, 2433 m asl, Pinus contorta, Pseudotsuga menziesii, Abies concolor, A. magnifica, etc., 16 Nov. 2011, D. Bojantchev DBB46740 (UC), 14 Nov. 2010, D. Bojantchev DBB38211 (UC). Additional specimens. Canada, British Columbia, VMS13, GenBank FJ717511 as ‘Cortinarius sp.’ (UBC). – USA, Oregon, Clackamas Co., Bull Run Watershed, Tsuga heterophylla, Pseudotsuga menziesii, 11 Nov. 1995, M.M. 95/500/ J.F. Ammirati 11701, GenBank EU057024 as ‘sp.’; Washington, Chelan Co., Chatter Creek, Tsuga heterophylla, Pseudotsuga menziesii, 6 Oct. 2007, J. Birkebak JMB10-06-2007-03 (UBC), GenBank HM068562 as ‘C. cupreorufus’. Notes — ITS sequence analysis shows C. luteicolor as a well-delimited species within clade /Laeticolores. Morphologically, it is not similar to any of the species we have studied. Initially, the species was described as a form of C. orichalceus var. olympianus (= C. pseudocupreorufus (A.H. Sm.) Niskanen, Liimat. & Ammirati). In the ITS regions it, however, differs from C. pseudocupreorufus by more than 25 substitutions and indel positions. Furthermore, Smith (1944) noticed several differences among these taxa: “This form (= C. luteicolor) differs from the typical form (= C. pseudocupreorufus) in lacking the faint lilac tint in the apex of stipe, in the pileus not becoming dark vinaceous red but instead merely dull cinnamon brown when fresh, and in the brighter yellow lamellae. The pilei of the herbarium specimens are also paler, but the bulb is deep purplish as in typical material of the variety (= C. pseudocupreorufus)”. Based on morphology and molecular data we conclude that C. luteicolor should be treated as a species. Description of the species is presented in Smith (1944). Typical for the species are at first rich dull yellow or yellow tinged pileus gradually becoming dull cinnamon, pale yellow gills, and subalmondshaped spores (9–11.5 × 6–7 µm). Cortinarius luteicolor grows in coniferous forests (Pseudotsuga, Tsuga) and is currently known from Pacific northwest of North America, from California and Washington, USA, and British Columbia, Canada. c b e Fig. 4 Spores of: a. C. flavipallens IK08-1729; b. C. boreicyanites CFP931; c. C. boreidionysae IK97-1220; d. C. cruentipellis IK98-2503; e. C. luteiaureus IK07-247a, in Melzer’s reagent. — Drawn by I. Kytövuori and T. Niskanen. — Scale bar = 10 µm. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium Cortinarius pseudocupreorufus (A.H. Sm.) Niskanen, Liimat. & Ammirati, stat. nov. & nom. nov. — MycoBank MB805886 Etymology. The name refers to the affinity to C. cupreorufus. Basionym. Cortinarius orichalceus var. olympianus A.H. Sm., Lloydia 7, 3: 184. 1944. Type. USA, Washington, Olympic National Park, Olympic Hot Springs, under conifers, 2 Oct. 1941, A.H. Smith 17513, barcode 10390 (holotype MICH). GenBank KF732367. Notes — In our phylogenetic analysis C. pseudocupreoru­ fus is placed as a sister species of C. cupreorufus Brandrud from which it differs by 10 substitutions and indel positions in ITS regions. The spores of C. pseudocupreorufus are 9–11 × 5 – 6.5 µm and somewhat amygdaloid, while the spores of C. cupreorufus are broader, 10 –11.5 × 6.5 –7.5 µm, and amygdaloid-citriform. Based on morphology and molecular data we conclude that C. pseudocupreorufus should be treated as a species. Description of the species is presented in Smith (1944). The species is so far only known from the type locality. /CyANITES Cortinarius boreicyanites Kytöv., Liimat., Niskanen & A.F.S. Taylor, sp. nov. — MycoBank MB805870; Fig. 4b Etymology. The name refers to the affinity with C. cyanites and distribution in boreal zone. Holotype. Sweden, Jämtland, Ragunda sn, Ragunda, Böle, at the riverside, in birch forest on rich soil (Betula, Picea), 24 July 1990, Brandrud et al. CFP931 (S). GenBank KF732296. Pileus 4–10 cm broad, hemispherical to convex, then expanded, distinctly innately fibrillose, bluish grey when young, later pale greyish brown. Lamellae emarginate, almost crowded, greyish blue when young, later brownish violet. Stipe 5–10 cm long, 1–2 cm thick at apex, 2.5–4 cm wide at base, clavate to bulbose, greyish blue. Universal veil pale greyish brown, abundant, forming girdles on stipe. Context violet when young, later in pileus and bulb white to brownish white, marbled hygrophanous, turning vinaceous red on exposure. Odour indistinct. Exsiccata: pileus reddish brown at the centre, greyish brown at the margin, stipe pale bluish greyish, brown at the base. In MLZ: Spores 9.1–9.8–10.4 × 5.4–5.8–6.3 µm, av. = 9.7–9.9 × 5.6–5.9 µm, Q = 1.58–1.69–1.80, Qav. = 1.66–1.73 (3 specimens, 120 spores, Fig. 4b), amygdaloid, with a blunt apex, fairly finely to moderately verrucose, many with few small golden yellow guttules, slightly dextrinoid. Basidia 36–45 × 7–9 µm (60 basidia), 4-spored, narrowly clavate, with a long, narrow pedicel, with blood red guttules, commonly with yellow foamy contents. Lamellar trama hyphae 4–10 µm wide, colourless to yellowish brown, smooth, with abundant small to large wormlike blood red guttules. Stipe apex hyphae 3–8 µm wide, pale yellowish, smooth, with small to large blood red guttules, the outermost ones c. 3 µm wide, ochraceous brown, mostly not guttulate. Pileipellis: epicutis very weakly gelatinous, uppermost hyphae 4–8(–13) µm wide, ochraceous brown, very thinly spotlike incrusted, not or weakly guttulate with small, golden yellow guttules, lower hyphae up to 10 µm wide, brown, smooth, with abundant small to large worm-like, blood red guttules. Hypoderm not developed. Ecology & Distribution — In boreal mixed forests of Picea, Betula and Populus in northern Europe. In Scotland also collected with Helianthemum. Fruiting in autumn. Other specimens examined. Finland, Pohjois-Savo, Kuopio, Vehmersalmi, submesic, mixed forest (Betula, Picea, Pinus), in grassy places, 1 Aug. 2003, T. Niskanen et al. 03-112 (H). – Great Britain, Scotland, Grampian, Braemar, Morrone Wood, with Helianthemum on rich calcareous soil, 8 Aug. 2010, A. Taylor 2010203 (UPS). 127 Notes — ITS sequence analysis shows C. boreicyanites as a well-delimited species within clade /Cyanites (Fig. 1). The ITS sequences of the species are identical and differ by more than 30 substitutions and indel positions from those of C. cyanites Fr. and C. violaceorubens Moënne-Locc. & Reumaux. Typical for C. boreicyanites are a pale greyish brown pileus, exsiccata with reddish brown pileus at the centre, greyish brown at the margin, and spores on average 9.7–9.9 × 5.6–5.9 µm. The species is so far only known from the middle boreal zone of Sweden, Finland and Scotland. Cortinarius cyanites has a more southern distribution extending from the hemiboreal zone of Finland and Sweden to France. In addition, the pileus is more bluish. Cortinarius violaceorubens has a more brownish pileus, dark dirty violaceous brown exsiccata, and larger spores (av. 9.9–11.0 × 6.3–6.6 µm). It grows in Picea dominated forests, often on rich soil. /DIONySAE s.l. Cortinarius boreidionysae Kytöv., Liimat., Niskanen & Dima, sp. nov. — MycoBank MB805894; Fig. 2e, 4c Etymology. The name refers to the affinity with C. dionysae and the northern distribution. Type. Finland, Perä-Pohjanmaa, Tervola, Peura, Raemäki, grass-herbspruce forest with spring-fed depressions, calcareous soil, 11 Sept. 1997, I. Kytövuori 97-1220 (holotype H; isotype NY). GenBank KF732488. Pileus 5–10 cm broad, hemispherical to convex, then expanded, glutinous, innately fibrillose, mustard brown to cocoa brown with a somewhat silky shining centre and olive yellowish tint on the margin when young. Lamellae emarginate, crowded, violaceous when young, later violet grey to pale brownish grey. Stipe 3–9 cm long, 1–1.8 cm thick at apex, 2–2.5 cm wide at base, with a marginate bulb, when young pale violaceous, becoming yellowish. Universal veil yellow at bulb margin. Con­ text in pileus white, in stipe violaceous, in bulb at first whitish, later brownish yellowish. Odour faintly farinaceous. Exsiccata: pileus uniformly orange brown, sometimes with a faint greyish tint, stipe somewhat paler. In MLZ: Spores 8.4–9.3–10.2 × 5.2–5.8– 6.1 µm, av. = 8.9–9.5 × 5.3–5.8 µm, Q = 1.54–1.67–1.80, Qav. = 1.63–1.67 (5 specimens, 160 spores, Fig. 4c), strongly citriform, strongly beaked, moderately, sharply verrucose, often with small coloured guttules, faintly to moderately dextrinoid. Basidia 23–32 × 7–9 µm (60 basidia), 4-spored, clavate, some with brownish yellow, foamy contents. Lamellar trama hyphae yellow, granuloseguttulate, blood red guttules absent. Stipe apex hyphae yellowish to yellow, smooth, with golden yellow small drops and large, blood red, worm-like guttules abundant in the outermost hyphae. Pileipellis: epicutis strongly gelatinous, uppermost hyphae 3–11 µm wide, mostly densely, spirally incrusted with small to large blood red guttules, lower down with a thick layer of somewhat wider hyphae filled with small to large or very large, worm-like, foamy, blood red guttules. Hypoderm well developed, yellow to red brownish. Ecology & Distribution — In northern boreal Picea abies dominated forests on calcareous soil, rare. Fruiting in autumn. Other specimens examined. Finland, Oulun Pohjanmaa, Kiiminki, Juuvansydänmaa, S part of Iso Juuvankangas, W of the lake Iso Juuvanjärvi, grass-herb-spruce forest with some Betula, Populus tremula and Pinus, on calcareous soil, 17 Aug. 2007, M. Toivonen & I. Kytövuori 07-245, H6000476 (H); Perä-Pohjanmaa, Tervola, Peura, Raemäki, grass-herb-spruce forest with spring-fed depressions, calcareous soil, 21 Aug. 1998, I. Kytövuori 98-1316, H6035751 (H), 21 Aug. 2007, M. Toivonen & I. Kytövuori 07-491, H6000724 (H); church village, Ala-Kirvesmaa, rich grass-herb-spruce forest with some Betula, Populus and Pinus, on calcareous soil, 24 Aug. 2007, I. Kytövuori 07-768, H6001001 (H); Petäjämaa, Kivimaa, 6 Sept. 2012, I. Kytövuori (H); Tornio, Korkiamaa, Runteli, rich grass-herb-spruce forest 128 with hardwood bushes and some pines, slightly paludified depressions, calcareous soil, 12 Aug. 1995, I. Kytövuori 95-210, H6035732 (H), 10 Sept. 1997, I. Kytövuori 97-1170 (H), 20 Aug. 1998, I. Kytövuori 98-1279, H6035740 (H), 22 Aug. 2007, I. Kytövuori 07-626, H6000859 (H); Kuusamo, Oulanka National Park, Ampumavaara, mesic to damp, herb-rich Picea abies forest, with some Pinus, Populus and Betula, 19 Sept. 2002, T. Niskanen et al. 02-876, H6033134 (H). Notes — Cortinarius boreidionysae reminds one of C. oli­ vaceodionysae A. Ortega, Vila & Fdez.-Brime, but the latter has paler, olive brown to yellowish brown pileus, somewhat larger, less distinctly and less regularly citriform spores, and a more southern distribution (up to the hemiboreal zone). The ITS sequences of C. boreidionysae are identical and it formed a well-supported clade in our phylogenetic analysis. It belongs to /Dionysae but further relationships were not solved. Based on ITS sequence comparison the closest species is C. olivaceodionysae from which it differs in ITS regions by 9 substitutions and indel positions, although two of them include intragenomic base polymorphisms. Cortinarius olivaceodionysae has been called C. dionysae Rob. Henry in northern Europe. Cortinarius dionysae, however, has been described from France (Henry 1933) as a species with a distinctly farinaceous smell and collected under Fagus – both characters in contradiction with our material of C. olivaceodionysae. To confirm the identity of C. dionysae we tried to sequence the type material but unfortunately we did not succeed. In our phylogenetic analysis all the taxa with a strong farinaceous smell, C. dionysae sensu Frøslev & Garnica, C. dionysae var. avellanus Rob. Henry ex Bidaud & Carteret and C. palazonianus Vila, A. Ortega & Fdez.-Brime, formed a well-supported subclade (1.00 PP) inside the clade /Dionysae. Of these, C. dionysae sensu Frøslev & Garnica and C. dionysae var. avellanus are potential candidates for the real C. dionysae. The former is collected under Fagus, but is so far only recorded from Germany and the spores are smaller than given in the original description. The latter is found in mixed forest in France but the spores fit well to the description. Further studies with French material are needed to confirm the identity of C. dionysae. Cortinarius cruentipellis Kytöv., Liimat., Niskanen & Dima, sp. nov. — MycoBank MB805872; Fig. 2f, 4d Etymology. The name refers to the large and abundant blood red drops/ guttules in the pileipellis. = Cortinarius luteoimmarginatus Rob. Henry sensu Jeppesen et al. (2012). = Cortinarius polymorphus Rob. Henry s.auct. p.p. Type. Sweden, Öland, Långlöt, Åstad, Nitares hägn, grassy pasture with Corylus and Juniperus, 13 Sept. 2003, I. Kytövuori, K. Liimatainen & T. Nis­ kanen 03-1451 (holotype H; isotype NY). GenBank KF732539. Pileus 3 –7.5 cm broad, hemispherical to convex, then expanded, unevenly wavy, olivaceous yellowish brown at centre, olivaceous to ochraceous yellow towards margin. Lamellae emarginate, almost crowded, pale grey when young, later pale greyish brown. Stipe 3–5 cm long, 0.5–1.4 cm thick at apex, 1.5–2.5 cm wide at base, with a marginate bulb, when young whitish, becoming slightly yellow with age. Universal veil ochraceous yellow at bulb margin. Context whitish. Odour indistinct. Exsiccata: pileus orange brown to dirty red brown at the centre, yellowish brown at the margin, stipe concolorous with the centre. In MLZ: Spores 9.5–10.4 –11.6 × 5.7–6.1–6.6 µm, av. = 10.3– 10.5 × 6.0 – 6.2 µm, Q= 1.59 –1.71–1.85, Qav. = 1.69 –1.73 (5 specimens, 160 spores, Fig. 4d), often strongly citriform, beaked, moderately, sharply verrucose, fairly faintly to moderately dextrinoid. Basidia 23 – 34 × 7.5 – 9.5 µm (50 basidia), 4-spored, clavate, some with yellow foamy contents. Lamellar trama hyphae yellowish, many with greenish yellowish, oily Persoonia – Volume 33, 2014 guttules. Blood red guttules absent. Stipe apex hyphae pale yellowish to greyish brownish to reddish brownish, smooth to finely or more strongly incrusted, somewhat granulose-guttulate, large blood red and smaller golden yellow guttules present in the outermost hyphae. Pileipellis: epicutis distinctly gelatinous, uppermost hyphae 4–10 µm wide, thin-walled, finely densely, spirally incrusted, mostly not guttulate, below these a thick layer of smooth to somewhat incrusted hyphae with abundant very long sausage-like guttules with blood red foamy contents. Hypoderm present, yellow brownish. Ecology & Distribution — Very rare in temperate to hemiboreal grass-herb forests with deciduous trees, mostly Corylus and Quercus, on calcareous soil. In addition, it occurs in halfopen deciduous vegetation in wooded pastures and parks. In Northwest Europe it is known in Denmark, Norway, Sweden and Estonia. Fruits in autumn. Other specimens examined. eStonia, Hiiumaa, Pühalepa, Sarve, dryish Corylus forest with Juniperus, Quercus and Populus, stony calcareous soil, 16 Sept. 2001, I. Kytövuori & T. Niskanen (Kytövuori 01-055, H). – Norway. Oslo, Bygdøy, Dronginberget south, deciduous forest, 26 Sept. 2004, T.E. Brandrud (Niskanen F04-983) (H7017763); Akershus, Asker, Spirodden, rich calcareous forest, Corylus, 8 Sept. 2011, I. Kytövuori 11-008 (H). – Sweden, Öland, Algutsrum, Gråbor Fornborg, rich grass-herb forest with Corylus, Quercus, Populus tremula and Picea, on sedimentary limestone, 30 Sept. 1988, I. Kytövuori 98-2503 (H); Öland, Torslunda, 1.5 km S from Borg, Corylus forest with some Picea, Betula and Quercus, 12 Sept. 2003, T. Niskanen D03-1393 & I. Kytövuori & K. Liimatainen, H7018687 (H). Notes — Cortinarius cruentipellis is a rather small, olivaceous tinted Phlegmacium of deciduous forests with citriform spores and strongly blood red pileipellis in MLZ. Of the other deciduous forest species with red MLZ reaction in pileus cuticle, C. gracilior (M.M. Moser) M.M. Moser is small and yellowish and has exsiccata with pale stramineous pileus and whitish stipe, C. leonicolor Reumaux (= C. anserinus (Velen.) Rob. Henry s.auct.) has much larger spores, and C. subdecolorans Langl. & Reumaux (= C. multiformium Cons. & Moënne-Locc.) pileipellis hyphae filled by mostly small guttules. Based on the ITS sequence analysis C. cruentipellis is a well-supported species (1.00 PP) belonging to the clade /Dionysae (0.82 PP). It has no close known relatives, and in the ITS regions it differs by more than 30 substitutions and indel positions from the closest species. Jeppesen et al. (2012) used the name C. luteoimmarginatus Rob. Henry for this species. In the original description of C. luteoimmarginatus (Henry 1939, as C. multiformis var. luteo­ immarginatus) the species is mentioned to have a clavate stipe, like the one of C. cliduchus, and this is also illustrated by Henry in Bidaud et al. (2012) whereas our species has a marginate bulb (Jeppesen et al. 2012). In addition, the spores are 11–11.5 × 5.5–6.5 µm, generally longer than those of our species and not overlapping with the average size of the spores (10.4 × 6.1 µm) or with the measurements of Jeppesen et al. (2012) (10–11 × 5.5–6.5 µm). In the original description no type has been designated but later Henry (1985) suggested a type (no. 1006) and a heterotype (no. 1014) for the species. The type of C. luteoimmarginatus, however, was not located in Paris (PC). A collection numbered as 1014 was found, but it was labelled as C. privignofulvus and represented a species of subg. Telamonia. Since the original description of C. luteoimmarginatus does not fit to our species we here describe it as new. /gLAUCOpODES Cortinarius subrubrovelatus (Bidaud) Kytöv., Liimat., Niskanen & Dima, comb. nov. — MycoBank MB805884 Basionym. Cortinarius glaucopus var. subrubrovelatus Bidaud, Atlas des Cortinaires 17, 2: 1237. 2008. Holotype. France, Ain, Farges, in mixed forest of Populus, Corylus, Fagus and Abies, 1 Nov. 1997, A. Bidaud 97-11-431 (PC). GenBank KF732317. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium Other specimens examined. Finland, Varsinais-Suomi, Lohja, Jalassaari, herb-rich Picea abies forest with some Corylus avellana, 19 Sept. 2004, I. Kytö­ vuori & T. Niskanen 04-881, H6031330 (H); Paimio, Huso, Kalkkimäki, below the old limestone quarries, moist spruce grass-herb forest with hardwood bushes, 6 Oct. 1985, I. Kytövuori 85-1553 (H). – Germany, Bayern, Oberjoch, conifer forest with Picea abies, 4 Oct. 2001, S. Garnica 4498 (TUB 011414), GenBank AY174787. Additional specimens. EStonia, Saare, Lümanda, in garden, with spruce and pine, V. Liiv 2009-10-29 (as ‘C. pini ’ TU106663/UDB011262). Notes — ITS sequence analysis shows C. subrubrovelatus as a well-defined species (1.00 PP) in /Glaucopodes. Typical for C. subrubrovelatus is pale brown to red brown pileus, bluish lamellae, and small, relatively broad spores (7.0–7.9 –8.8 × 4.5–5.0–5.4 µm, av. = 7.7–8.0 × 4.8–5.2 µm, Q = 1.44–1.56– 1.70, Qav. = 1.52–1.61 (3 specimens, 180 spores)). From its closest relatives C. subfoetens and C. glaucopus it differs by 2 and 4 substitutions and/or indel positions. Cortinarius sub­ foetens has larger, more fusoid and narrower, and less dextrinoid (8.2– 8.9 –9.7 × 5.0– 5.3 –5.4 µm, Q = 1.58–1.69 –1.82) spores, and the spores of C. glaucopus are relatively narrower as seen in the Q-values (7.3 – 8.1– 9.1 × 4.5 – 5.0 – 5.4 µm, Q = 1.54 –1.64 –1.82). The latter also has somewhat more reddish brown colours deep in the pileipellis and less incrusted upper hyphae than those of C. subrubrovelatus. The differences in the ITS region between the three species are not large but since all three groups are represented by several collections and also have morphological differences we conclude that C. subrubrovelatus should be treated as a species. Cortinarius subrubrovelatus grows in coniferous forests on calcareous soil and is known from Central Europe and hemiboreal zone of northern Europe. /ELASTICI & pERCOMES Cortinarius luteiaureus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805895; Fig. 4e Etymology. The name refers to the colour of the pileus. Type. Finland, Oulun Pohjanmaa, Kiiminki, Juuvansydänmaa, S part of Iso Juuvankangas, W of the lake Iso Juuvanjärvi, grass-herb Picea abies forest with some Betula, Populus tremula and Pinus, on calcareous soil, 17 Aug. 2007, M. Toivonen & I. Kytövuori 07-247b, H6033617 (holotype H; isotype NY). GenBank KF732568. Pileus 4–7 cm broad, convex, then plano-convex, with a low and broad umbo, viscid to glutinous, not fibrillose, yellow to brownish yellow. Lamellae emarginate, almost crowded, greyish white, later very pale greyish brown. Stipe 5–10 cm long, 1–1.5 cm thick at apex, 1.5–2 cm wide at base, almost cylindrical with clavate to subbulbous base, white. Universal veil yellow, forming girdles on stipe. Context white. Odour not recorded. Exsiccata: pileus yellow brownish, stipe whitish. In MLZ: Spores 9.7–10.6 –11.6 × 5.7–6.1–6.6 µm, Q = 1.62– 1.74–1.92 (1 specimen, 60 spores, Fig. 4e), narrowly amygdaloid, with rounded apex, moderate to fairly strongly verrucose, moderately dextrinoid, often with dark red brown angular granules in the spore. Basidia 32–44 × 8–10 µm (20 basidia), 4-spored, clavate, sand brown, with fairly small granules and chips. Lamellar trama hyphae with moderate to very large sand brown to red brown granules. Stipe apex hyphae sand brown, densely granulose, outermost ones more orange or reddish, not granulose. Pileipellis: epicutis strongly gelatinous, uppermost hyphae 2–4 µm wide, ochraceous yellow to ochraceous brown, mostly not granulose, lower down 4–10 µm wide, full of small to very large dark red brown granules. Hypoderm absent. Ecology & Distribution — In coniferous forest (Picea) on calcareous soil. Known from northern Finland, Oulun Pohjanmaa. No sequences of this species exist in public databases. 129 Notes — Cortinarius luteiaureus resembles somewhat a species which we assume could be called C. rhizophorus Bidaud & Cons. but we have not studied the type material of the species yet. Cortinarius cf. rhizophorus is larger, somewhat paler, has larger spores (10.2–11.3–12.5 × 6.1–6.5–7.0 µm) and occurs in temperate to hemiboreal broad-leaved and coniferous forests. Cortinarius varius (Schaeff.: Fr.) Fr. is also somewhat similar but has blue lamellae, stout, clavate stipe, wider spores, and hypoderm in the pileipellis. Cortinarius luteiaureus differs in ITS regions by more than 30 substitutions and indel positions from the closest species found with the BLAST. Based on our phylogenetic analysis it belongs to the large /Elastici & Percomes clade. /INFRACTI Cortinarius infractiflavus (M.M. Moser) Kytöv., Niskanen, Liimat., Bojantchev & Ammirati, stat. nov. & nom. nov. — MycoBank MB805887 Etymology. The name refers to the affinity to C. infractus and the yellowish pileus. Basionym. Cortinarius infractus var. flavus M.M. Moser, Mycotaxon 72: 313. 1999. Type. USA, Wyoming, Shoshone Natl. Forest, prope Brooks Lake Lodge, in subalpine coniferous forest (Picea engelmanii, Abies lasiocarpa), 12 Aug. 1997, M. Moser 97/169 (holotype IB). GenBank KF732327. Other specimens examined. Bulgaria, Pirin Mt, Bansko locality, 1600 m asl, under Picea abies, 7 Oct. 2009, D. Bojantchev, DBB19634 (UC). – Finland, Sompion Lappi, Pelkosenniemi, Suvanto, Kalkkivaara, half-open dry Pinus sylvestris forest with Picea, Juniperus, Betula, Populus and Alnus, dolomite rock, 26 Aug. 1992, I. Kytövuori 92-1109 (H). Additional specimen. Canada, British Columbia SMI286, GenBank FJ039612 (UBC). Notes — Cortinarius infractiflavus formed a well-supported clade (1.00 PP) in our phylogenetic analysis. It belongs to sect. Infracti and differs from the sister species C. infractus by 9 substitutions and indel positions in ITS regions. Morphologically, it can be distinguished from C. infractus by the yellowish colours of the pileus, paler gills, a nearly mild taste and larger spores (7.5– 8.2–9.1 × 6.3–6.7–7.0 µm, Q = 1.16–1.23 –1.30; C. in­ fractus 7.3– 8.0–8.8 × 5.7– 6.1–6.6 µm, Q= 1.23–1.30 –1.38). Cortinarius infractiflavus grows in boreal and mountainous conifer forests (Picea, Abies) and is widespread occurring from Wyoming, western USA to Finland and Bulgaria. Full description of the species can be found from Moser & Ammirati (1999). /MULTIFORMES Cortinarius caesiolamellatus (Bidaud) Kytöv., Liimat., Niskanen, Brandrud, Frøslev & A.F.S. Taylor, comb. nov. — MycoBank MB805885 Basionym. Cortinarius rufoallutus var. caesiolamellatus Bidaud, Atlas des Cortinaires 16: 1095. 2006. Type. France, Ain, Bugey, col de Bérentin, in young Picea abies plantation, on calcareous soil, 3 Oct. 1993, A. Bidaud PML4905 (holotype PC). GenBank KF732414. = Cortinarius multiformis var. caesiophyllus Moënne-Locc., Atlas des Cortinaires 16: 1095. 2006. — Type. P. Moënne­Loccoz PML882 (holotype PC), France, Savoie, Arith, under Picea on calcareous soil, 3 June 1988. GenBank KF732351. Illustrations — Bidaud et al. (2006: pl. 581, 587). Pileus 4–8 cm broad, hemispherical to plano-convex, sometimes radially rugulose towards the margin, viscid, red brown to ochraceous brown, rarely bluish brown, becoming paler ochra- 130 ceous brown with age, often bi-coloured, outer part hygrophanous and darker (grey) brown. Lamellae emarginate, almost crowded, pale grey with a bluish tint when young, later pale brown. Stipe 4–8 cm long, 0.7–1.7 cm thick at apex, cylindrical, slender, 1.5–2.5 cm wide at base, mostly bulbous but the bulb often fairly small, fibrillose (not glossy like C. multiformis), whitish but often 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 fairly thin, whitish, brownish below the pileus cuticle, in the stipe with a bluish tint or not when young. Odour faint to distinct of honey in the context of the stipe base. Exsiccata: pileus greyish to reddish brown, stipe very pale. In MLZ: Spores 8.2–9.3–10.7 × 5.0–5.6–6.3 µm, av. = 8.5–9.9 × 5.2–5.9 µm, Q = 1.51–1.65–1.85, Qav. = 1.57–1.74 (7 specimens, 240 spores), ovoid-amygdaloid to narrowly ovoid-ellipsoid, with a fairly long, blunt, tapering apex, moderately to strongly verrucose, warts wide, anastomosing, slightly to fairly strongly dextrinoid. Basidia 25–34 × 7.5–9 µm (100 basidia), 4-spored, clavate, mostly with fairly wide base. Many basidia and subhymenium with yellowish, granulose/guttulate contents. Pilei­ pellis: epicutis with a fairly thick, gelatinous layer with some 2–3 µm wide, erect-entangled, smooth, obscure, colourless hyphae. Hypoderm present, fairly thin-walled, with pale brownish parietal pigment, some small brown spots present, large yellow to yellow brown spots absent, few spirally incrusted hyphae present deep in the pileipellis. Ecology & Distribution — In montane to middle boreal, mesic coniferous forests, often on rich soil. In Europe apparently mainly/ only under Picea, but in USA also found under Pinus. Known from Central and northern Europe, and Washington, USA, and considered occasional. The fruiting period is very long, once collected in June (France) and once in November (USA). Other specimens examined. EStonia, Saaremaa, Mustjala, Võhma, dryish pine heath forest with some spruce, on calcareous soil, 16 Sept. 1993, I. Kytövuori 93-1336 (H). – Finland, Uusimaa, Kunnarla, Myllyoja, grass-herb forest with some pines and hardwood trees, 22 Sept. 1994, I. Kytövuori 94-852 (H); Pohjois-Savo, Kuopio, Vehmersalmi, Putroniemi, Roikanselän rantaMäkijärvi E, under Picea, 30 years old forest, 26 Sept. 2009, J. Ruotsalainen 8103F, H6011464 (H). – France, Oyonnax SE, Giron, Forêt de Champfromier, rich conifer forest with Abies alba, Picea abies and Fagus, 27 Oct. 1994, P. & I. Kytövuori 94-1897 (H); Savoie, Arith, under Picea on calcareous soil, 3 June 1988, P. Moënne­Loccoz PML882 (PC). – Germany, Baden-Württemberg, Thölendorf, Picea, 11 Oct. 1998, UL 98/122 (TUB 011841), GenBank AY669531 as ‘C. allutus’; Schwaben, Ehingen a.d. Donau, Kohlhau, Picea plantation on calcareous soil, 28 Sept. 2010, G. Schmidt­Stohn & T.E. Bran­ drud, TEB 428-10 (O, TUB). – Norway, Oslo, Steinbruvannet-Røverkollen, Picea forest of somewhat richer, low-herb type, 4 Sept. 2011, T.E. Brandrud 687-11 (O). – Sweden, Uppland, Uppsala, Nåsten, mixed forest, 27 Aug. 2005, A. Taylor 2005085, UNITE No. UDB002202 as ‘C. allutus’ (UPS); Vänge, Fiby urskog, virgin spruce forest with Pinus, Betula, Populus tremula, Colylus, Alnus glutinosa, etc., with fairly rich depressions, decaying logs very abundant, 8 Sept. 1994, P. & I. Kytövuori 94-309 (H). – USA, Washington, Grays Harbor Co., Ocean Shore State Park, under Pinus on sandy soil, 11 Nov. 2009, J.F. Ammirati & T. Niskanen 09-201 (H). Notes — Cortinarius caesiolamellatus and C. caesiophyl­ loides are easily distinguished from the related species in /Multiformes by their initially bluish tinged lamellae and often also stipe apex. Based on material seen so far and available descriptions, these two blue-gilled species are hardly distinguishable macromorphologically, but C. caesiolamellatus differs in slightly larger and more strongly verrucose spores. Furthermore, there seems to be a geographical differentiation in Europe; C. caesiolamel­ latus apparently being mainly a Central European species, whereas C. caesiophylloides is hitherto found only in northern Europe. If bluish tints have gone, C. caesiolamellatus can also be confused with conifer associated C. rufoallutus Rob. Henry ex Bidaud & Reumaux, C. multiformis Fr. and C. talimultiformis Kytöv., Liimat., Niskanen, A.F.S Taylor & Sesli. The latter two, Persoonia – Volume 33, 2014 however, have less strongly ornamented spores and lack the yellow to yellow brown large spots deep in the pileipellis. Cor­ tinarius rufoallutus differs from C. caesiolamellatus by stouter appearance, small amygdaloid-fusoid, finely verrucose spores and abundant strongly spirally incrusted red brown hyphae deep in the pileipellis. In our phylogenetic analysis C. caesiolamellatus forms a wellsupported group (1.00 PP) and belongs together with C. cae­ siophylloides and C. pallidirimosus in a strongly supported (1.00 PP) subclade in clade /Multiformes (1.00 PP). The species was originally described twice in Bidaud et al. (2006), as a variety of both C. rufoallutus (var. caesiolamellatus) and C. multiformis (var. caesiophyllus). Based on the molecular and morphological data neither of those relationships is supported. Cortinarius caesiolamellatus clearly represents a distinct species and is here combined at species level. Cortinarius caesiophylloides Kytöv., Liimat., Niskanen, Brandrud & Frøslev, sp. nov. — MycoBank MB805873; Fig. 2g, 5a Etymology. The name refers to the bluish tints in lamellae and affinity with C. caesiolamellatus (= Cortinarius multiformis var. caesiophyllus). Type. Finland, Etelä-Savo, Joutsa, Koivuranta, W of Rakkolanselkä, fairly young, mesic to damp, Picea abies-dominated forest with some Betula and Pinus sylvestris, 30 Aug. 2005, T. Niskanen et al. 05-016, H6029792 (holotype H; isotype NY). GenBank KF732572. Pileus 4–8 cm broad, hemispherical to plano-convex, viscid, apricot to redbrown-coloured, with hygrophanous streaks or outer zone, hygrophanous areas somewhat darker umber to less vivid (grey) brown, becoming paler ochraceous brown to ochraceous yellow with age, sometimes almost whitish ochre at centre. Lamellae emarginate, almost crowded, pale grey with a bluish tint when young, later pale brown. Stipe 5–11 cm long, 1–1.5 cm thick at apex, 1.5–3.5 cm wide at base, with a more or less distinct marginate bulb, whitish but usually with a bluish tint at apex when very young, often becoming pale brown with age, but not as pronounced as with C. multiformis. Universal veil white, sparse to hardly visible, at bulb margin. Context white, slightly bluish at the apex of the stipe when young. Odour faint to distinct of honey in the context of the stipe base. Exsiccata: pileus pale dirty brown, stipe somewhat lighter. In MLZ: Spores 8.6–9.7–10.9 × 5.2–5.8–6.3 µm, av. = 9.0–10.2 × 5.5–6.0 µm, Q = 1.54–1.68–1.85, Qav. = 1.63–1.73 (4 specimens, 240 spores, Fig. 5a), amygdaloid, with fairly narrow apex, sometimes almost ellipsoid, finely to moderately verrucose, warts often rounded-confluent, slightly to fairly strongly dextrinoid; with a bit narrower apex, thinner wall and finer verrucosity than in C. multiformis. Basidia 24–37 × 7–9 µm (60 basidia), 4-spored, clavate, almost colourless. Lamellar trama hyphae very pale yellowish, smooth, concolorous guttulate. Stipe apex hyphae very pale yellow, smooth, somewhat concolorous guttulate. Pileipellis: epicutis with a clear gelatinous layer with sparse, erectentangled, 2–3 µm wide, smooth, colourless and very obscurely seen hyphae. Hypoderm present, hyphae fairly thin-walled, with pale yellowish brown amber-like parietal pigment, small brown encrust-spots present, large yellow brown spots absent, a few spirally incrusted (zebra-striped) hyphae seen deep in the cuticle. Ecology & Distribution — In mesic coniferous forests, presumably with Picea, mainly in richer low-herb types, sometimes on calcareous soil. Known only from Fennoscandia and considered occasional. Fruits from August to September. Other specimens examined. Finland, Kainuu, Paltamo, Oikarilankylä, Kivesvaara, old mesic spruce forest with some Betula, Pinus and Populus tremula, 11 Sept. 2008, I. Kytövuori 08-1554, H6032621 (H); Sompion Lappi, Pelkosenniemi, Suvanto, Kalkkivaara, steep SW slope, dry pine forest with Juniperus, Betula, Populus, Alnus incana, 26 Aug. 1992, I. Kytövuori 92-3044, H6032620 (H). – Norway, Nord-Trøndelag, Stjørdal, Beistadvollen, calcareous spruce forest (on karst surfaces), 13 Aug. 2009, T.E. Brandrud 277-09 (O). K. Liimatainen et al.: Identification and nomenclature of Phlegmacium a b 131 c d Fig. 5 Spores of: a. C. caesiophylloides TN05-016; b. C. melleicarneus IK01-053; c. C. pallidirimosus IK95-585; d. C. talimultiformis AT2004096, in Melzer’s reagent. — Drawn by I. Kytövuori and T. Niskanen. — Scale bar = 10 µm. Notes — Cortinarius caesiophylloides is a typical member of sect. Multiformes. Together with C. caesiolamellatus they are the only known species of the section with bluish tinted lamellae when young. Cortinarius caesiolamellatus differs from C. caesiophylloides by more strongly ornamented spores. When bluish tints are absent they can be confused with C. multi­ formis and C. talimultiformis, but in the latter two there are abundant, yellow to yellow-brown, large spots deep in the pileipellis. Those are practically lacking from C. caesiophylloides and C. caesiolamellatus. In C. multiformis the honey smell is either lacking or is very weak, and the stipe becomes strong brass brown with age. Cortinarius caesiophylloides is strongly supported (1.00 PP) in our phylogenetic analysis. It belongs to /Multiformes (1.00 PP) and further, in a well-supported (1.00 PP) subclade with C. caesiolamellatus and C. pallidirimosus. The ITS sequences of C. caesiophylloides have one base polymorphism and the maximum pairwise distance is zero. The difference to C. palli­ dirimosus is 11 substitutions and indel positions. Cortinarius melleicarneus Kytöv., Liimat., Niskanen & Brandrud, sp. nov. — MycoBank MB805874; Fig. 5b, 6a Etymology. The name refers to the colour of the pileus. Type. EStonia, Hiiumaa, Pühalepa, Sarve, Soonlepa, deciduous forest (Corylus, Quercus) with some Pinus on calcareous soil, 16 Sept. 2001, I. Kytövuori 01-053 (holotype H; isotype NY). GenBank KF732577. Pileus 4–10 cm broad, hemispherical to convex, with rather persistently incurved margin, then expanded, sometimes somewhat silvery-silky from fine veil remnants, cream-coloured, pale yellow brown, honey brown to more grey brown, with hygrophanous streaks or patches/zones near margin; hygrophanous zones somewhat darker grey brown, almost with an olivaceous brown tinge. Lamellae emarginate, crowded, first pale greyish white, later pale greyish brown. Stipe 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, short and robust, white. Universal veil very sparse at bulb margin, white. Context in pileus pale yellow brown, marbled hygrophanous flesh-coloured, in stipe white. Odour not recorded. In MLZ: Spores 7.9– 8.7–9.5 × 4.3–4.7–5.2 µm, av. = 8.6–8.9 × 4.7– 4.8 µm, Q = 1.67–1.86 – 2.00, Qav. = 1.85 –1.89 (2 specimens, 120 spores, Fig. 5b), amygdaloid to fusoid, with a low suprahilar depression and blunt apex, moderately to fairly strongly, densely and coarsely, somewhat unevenly, not sharply verrucose, slightly to moderately dextrinoid. Basidia 25–31 × 7–8 µm (40 basidia), 4-spored, clavate, many granulose-guttulate. Lamellar trama hyphae pale yellow, smooth. Stipe apex hyphae very pale yellowish, smooth, coloured guttules absent. Pileipellis: epicutis gelatinous, with 3–8 µm wide, thin-walled, smooth to very finely incrusted, colourless and very obscure hyphae. Hypoderm present, with pale yellowish brown walls, few incrusted hyphae and small brown spots deep in the pileipellis. Ecology & Distribution — With thermophilous deciduous trees (Corylus, Quercus) on calcareous soil, including nearshore sandy shell-beds. Known from Estonia and Norway. Other specimens examined. Norway, Aust-Agder, Grimstad, Fevik, under Quercus, Fagus and (planted) Larix also present, on sandy soil, rather rich, probably with shell-bed deposits, 21 Sept. 1994, leg. I­L. Fonneland, det. T.E. Brandrud 86-94, O 125960 (O, sub nom. C. areni­silvae). Notes — Cortinarius melleicarneus belongs to /Multiformes. It differs from the other species of the group by honey-coloured pileus, fusoid spores, and habitat with deciduous trees on calcareous soil. 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. Corti­ narius talus has furthermore usually an innately fibrillose pileus structure and rarely possess short-stemmed, compact carpophores. Cortinarius cruentipellis grows in similar habitats, but it has ochraceous yellow universal veil on bulb margin, larger spores, and red-colouring pileipellis in MLZ. In dry conditions, C. melleicarneus might resemble C. areni­silvae (Brandrud) Brandrud which also occurs in sandy soils, and the Norwegian collection was originally determined as C. areni­silvae (based on dry material and collectors notes). ITS sequences of the two C. melleicarneus specimens are identical and it forms a clade in our analysis. It differs by 11 substitutions and indel positions from C. talimultiformis, 14 from C. talus, 15 from C. multiformis, and 17 from C. rufoallutus Rob. Henry ex Bidaud & Reumaux. Cortinarius pallidirimosus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805875; Fig. 5c, 6b Etymology. The name refers to the pale pileus with streaks. Type. Finland, Inarin Lappi, Utsjoki, Kevo, Tsieskuljohka, mesic heath forest with Betula and Pinus, with some moist depressions, 17 Aug. 1995, I. Kytövuori 95-585, H6035694 (holotype H; isotype NY). GenBank KF732578. Pileus 3–9 cm broad, hemispherical to convex, then expanded, viscid, very finely innately fibrillose, whitish to cream-coloured, centre brownish yellow, becoming ochraceus with age, with hygrophanous streaks. Lamellae emarginate, crowded, pale greyish brown when young, later pale brown. Stipe 6–13 cm long, 0.7–1.5 cm thick at apex, 1.5–2.5 cm wide at base, clavate to almost cylindrical, whitish, becoming very pale brown with age. Universal veil white, sparse. Context white. Odour in context honey-like. Exsiccata: pileus cream-coloured to ochre brownish, somewhat darker at the centre, lighter towards the margin, stipe somewhat lighter than pileus. In MLZ: Spores 8.6–9.6–10.7 × 5.2–5.7–6.1 µm, av. = 9.0–10.1 × 5.5–5.9, Q = 1.54–1.70–1.85, Qav. = 1.60–1.76 (12 specimens, 320 spores, Fig. 5c), amygdaloid to amygdaloid-ellipsoid, moderately to fairly strongly, unevenly, sometimes coarsely verrucose, warts fairly wide but not high, slightly (to moderately) dextrinoid. Basidia 24–36 × 7.5–9 µm (100 basidia), 4-spored, clavate, almost colourless. Lamellar trama hyphae pale pellucid yellowish, smooth, somewhat concolorous granulose-guttulate. 132 Persoonia – Volume 33, 2014 Fig. 6 Photo of: a. C. melleicarneus IK01-053; b. C. pallidirimosus IK95-585; c. C. talimultiformis AT2004096; d. C. balteatialutaceus IK09-751; e. C. bal­ teatibulbosus IK98-1624; f. C. balteaticlavatus IK95-382; g. C. brunneiaurantius JV17979; h. C. caesiocolor IK00-029. — Photos: a, b, d–f, h. I. Kytövuori; c. A. Taylor; g. J. Vauras. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium Few small mounds of colourless crystals sometimes present in the lamellar trama. Stipe apex hyphae pale yellowish, smooth. Pileipellis: epicutis with a clear slime layer with 1.5–3 µm wide, smooth, colourless and very obscure hyphae. Hypoderm present, very pale brownish to almost colourless. Ecology & Distribution — In the middle to northern boreal, mesic, mixed forests with Betula, among mosses, on oligotrophic to eutrophic soil. Often solitarily or in small groups. Known from Fennoscandia, Sakha, Russia, and Oregon, USA, and considered occasional. Fruiting in late summer to autumn. Other specimens examined. Finland, Kainuu, Puolanka, Väyrylä, Körölä, grass-herb-spruce forest with some pines and hardwood bushes, 15 Sept. 1997, I. Kytövuori 97-1523, H6035715 (H); Perä-Pohjanmaa, Rovaniemi, Louevaara, Tuohilaki nature reserve area (west), eutrophic, submesic to mesic Picea abies forest with Betula, Populus tremula and some Pinus sylvestris, 29 Aug. 2004, T. Niskanen & K. Liimatainen 04-470, H6029374 (H); Tervola, Louepalo, Kätkävaaran marmorilouhos, E of the western quarry, S of the track, gently E sloping forest of young Betula and Picea, with some Populus tremula, Alnus incana and Salix spp., 23 Aug. 2007, I. Kytövuori 07-692, H6000925 (H); Peura, Kaitalampi, Kaitaharju, half-open, eutrophic grass-herb-spruce forest with some pines, calcareous soil, 5 Sept. 1992, I. Kytövuori 92-1779, H6035693 (H); Koillismaa, Kuusamo, Oulanka National Park, Ampumavaara, mesic to damp, herb-rich Picea abies forest with some Pinus, Populus and Betula, 19 Sept. 2002, T. Niskanen et al. 02-859, H6033129 (H); Kittilän Lappi, Kolari, Teuravuoma-Lappea (Korkealehdontie), Tappikumpu, mesic sprucebirch forest, 18 Aug. 1998, I. Kytövuori 98-1078, H6035705 (H); Sompion Lappi, Sodankylä, Tähtelä, Hassikankaannokka, mesic spruce forest with some pines and hardwood trees, 24 Aug. 1992, I. Kytövuori 92-966 (H); Inarin Lappi, Utsjoki, Kevo, Tsieskuljohka, mesic heath forest with Betula and Pinus, some moist depressions, 17 Aug. 1995, I. Kytövuori 92-540, H6035704 (H). – Norway, Troms, Storfjord, Skibotndalen, Lullesletta, dry pine forest with Betula and Salix, 13 Aug. 1998, I. Kytövuori 98-711 (H). – Sweden, Jämtland, Bodsjö, Sidsjö, Stuguberget, dryish, fairly young spruce-pine forest with some hardwood trees and bushes, some moist depressions, 4 Sept. 1997, I. Kytövuori 97-699, H7019576 (H); Frösö, Böle, Fillstabäcken, damp to mesic coniferous forest (Picea, Pinus) on calcareous soil, 2 Sept. 2003, Niskanen et al. 03-1058, H7018404 (H). Additional specimens. ruSSia, Sakha, Khangalasskiy Ulus, Myachei-Sise Mts, Larix gmelinii forest with Betula platyphylla, 8 Aug. 1999, U. Peintner IB19990590 (IB), UNITE No. UDB001073. – USA, Oregon, source: mycorrhizal root tip, clone=”8_73M8, GenBank JQ393042. Notes — Cortinarius pallidirimosus can easily be identified by the cream-coloured, hygrophanous streaked pileus, honeylike smell in context, large, amygdaloid to amygdaloid-ellipsoid spores, and habitat with Betula in the boreal zone. The other Betula associated species in sect. Multiformes, C. talus Fr., has relatively wider, ochraceous yellow pileus and smaller spores (7.3 – 8.0 – 8.8 × 4.5 – 5.0 – 5.2 µm). Cortinarius gentianeus Bidaud and C. cremeiamarescens Kytöv., Liimat. & Niskanen are reminiscent of C. pallidirimosus, but the former two have a bitter tasting pileus, indistinct odour, and smaller, citriform to narrowly fusoid spores. In addition, the lamellae and pileipellis of both species are strongly red in MLZ. The placement of C. pallidirimosus in /Multiformes is well supported (1.00 PP) and it forms a strongly supported subclade with C. caesiolamellatus and C. caesiophylloides. From its sister species C. caesiophylloides it differs by 11 substitutions and indel positions in ITS regions. The sequences of C. palli­ dirimosus have one base polymorphism and six intraspecific variation sites and the maximum pairwise distance is six. The amount of variation is highly compared to many other species in the genus Cortinarius in which the variation typically is 0–2 substitutions and/or indel positions. Most likely the data includes a close sister species, but more specimens would be needed to study this complex more in detail. Cortinarius talimultiformis Kytöv., Liimat., Niskanen, A.F.S. Taylor & Sesli, sp. nov. — MycoBank MB805876; Fig. 5d, 6c Etymology. The name refers to the appearance of the species, which has features from both C. multiformis and C. talus. 133 Type. Sweden, Uppsala, Hässelby Park, mixed forest, 11 July 2004, A. Taylor AT2004096 (holotype UPS; isotype S) UNITE No. UDB001167. GenBank KF732583. = Cortinarius aurantionapus var. similis Moënne-Locc., Atlas des Cortinaires 16: 1096. 2006. — Type. P. Moënne­Loccoz, PML883 (holotype PC), France, Haute-Savoie, Aviernoz, under Picea abies, on calcareous soil, 6 June 1988, . Illustration — Bidaud et al. (2006: pl. 596). Pileus 4–12 cm broad, hemispherical to plano-convex, viscid, whitish fibrillose, especially near the margin, ochraceous yellow to red brown, with strongly hygrophanous streaks. Lamellae emarginate, crowded, pale grey when young, later pale greyish brown. Stipe 4–10 cm long, 1–2 cm thick at apex, 2–3.5 cm wide at base, clavate or with a somewhat marginate bulb, white. Universal veil white, sparse, sometimes fairly abundant at pileus margin. Context white. Odour slightly honey-like at the context of the base of the stipe. Exsiccata: pileus leather brown to yellow brown to mahogany brown, sometimes whitish variegate at the centre, stipe whitish to pale brownish. In MLZ: Spores 8.4–9.5–10.7 × 5.0–5.5–6.1 µm, av. = 8.9–9.9 × 5.3–5.8 µm, Q = 1.58–1.73–1.87, Qav. = 1.68–1.78 (9 specimens, 320 spores, Fig. 5d), amygdaloid to narrowly amygdaloid to amygdaloid-fusoid, with a long, narrowish apex, rather weakly to fairly strongly, lowly verrucose, slightly to moderately dextrinoid. Basidia 27–36 × 7–9 µm (80 basidia), 4-spored, clavate. Lamellar trama hyphae very pale yellowish, smooth. Stipe apex hyphae 4–8 µm wide, pale yellowish, smooth, some outermost ones 2–3 µm wide, in entangled bundles. Pileipel­ lis: epicutis with fairly thick, clear slime layer with 1.5–2.5 µm wide, colourless, hardly visible hyphae. Hypoderm present, in its upper part large, brownish yellow pigment spots abundant. Ecology & Distribution — In hemiboreal to boreal, mesic coniferous forests with Picea and Abies. Known from North and Central Europe, and East Black Sea Region’s mountains in Turkey, considered common. Basidiocarps occur from June to late August. Other specimens examined. Finland, Pohjois-Häme, Virrat, Killinkoski, Abies sibirica forest, 30 Aug. 1966, I. Kytövuori, H6032747 (H); Oulun Pohjanmaa, Kiiminki, Keskikylä, Pikkuhalmeenmaa, grass-herb-spruce forest with some Betula, Populus tremula and Pinus, on calcareous soil, 15 Aug. 2007, M. Toivonen & I. Kytövuori 07-131, H6000360 (H); Perä-Pohjanmaa, Tornio, Kalkkimaa, Kalkkimaan lehto (NW), mainly grass-herb forest with Picea, Betula, Populus tremula, Juniperus and Pinus, on calcareous soil, 22 Aug. 2007, I. Kytövuori 07-605, H6000838 (H). – France, Haute-Savoie, Aviernoz, under Picea abies, on calcareous soil, 6 June 1988, P. Moënne­ Loccoz, PML883 (PC, holotype of C. aurantionapus var. similis). – Germany, Baden-Württemberg, Titisee, saurer Boden unter Fichten, 10 Sept. 2002, Saar 4855, TUB 011864 (TUB), GenBank AY669532 as ‘C. allutus’ (TUB). – Norway, Oppland, Gran, Lygna, mesic spruce forest with some Betula and Pinus, 13 Sept. 2010, E. Bendiksen & I. Kytövuori (H). – Turkey, East Black Sea Region, Trabzon, Macka, Gurgenagac village, Picea orientalis forest, 7 July 2010, E. Sesli, SES 2741. Notes — The appearance of C. talimultiformis is rather variable, but most often it is fairly low, wide and stout. The colour of the pileus varies from ochraceous yellow to mahogany brown and the form of the stipe from clavate to bulbous. Typical for the species, however, are the white fibrils on the pileus margin, narrow, prominently verrucose spores, abundant large yellow spots in the pileipellis, and habitat with Picea or Abies. The fruiting period of this species starts early and it can be found in early June. The sister species C. multiformis Fr. grows in similar habitats but has no fibrils on the pileus, the spores are moderately to strongly dextrinoid, fairly finely verrucose, and abundant large brownish yellow spots are present deep in the pileipellis. The colour of the pileus in the exsiccata is uniformly red brown. The other common species, C. talus Fr., is associated with deciduous trees, is pale ochraceous yellow, and the spores are smaller (7.3– 8.0 –8.8 × 4.5– 5.0 –5.2 µm). 134 Cortinarius talimultiformis formed a well-supported group in our phylogenetic analysis. It is a sister species of C. multiformis from which it differs by 6 substitutions and indel positions in ITS regions. /pHLEgMACIOIDES Cortinarius balteatialutaceus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805877; Fig. 6d, 7a Etymology. The name refers to the colour of the pileus. Type. Sweden, Jämtland, Frostviken, Jormlien, Säterklumpen, Betula forest with solitary Picea, 7 Sept. 2009, P. & I. Kytövuori 09-751 (holotype H; isotype NY). GenBank KF732586. Pileus 6–12 cm broad, hemispherical to plano-convex, with a fairly persistently involute margin, first very slightly viscid, dry with age, pale brown with a whitish to very pale brown margin, becoming darker brown from centre. Lamellae emarginate, crowded, pale brownish grey when young, later pale brown. Stipe 5–7 cm long, 1.3–2.5 cm thick at apex, 1.8–3.5 cm wide at base, clavate, sometimes with an almost marginate bulb, whitish. Universal veil white, sparse. Context white. Odour indistinct. KOH­reaction in context yellow. Exsiccata: pileus leather brown at centre, paler at margin, stipe very pale brownish. In MLZ: Spores 9.1–10.0 –10.9 × 5.0–5.4 –5.9 µm, av. = 9.8– 10.4 × 5.3–5.5 µm, Q = 1.75–1.85 –1.98, Qav. = 1.78–1.87 (5 specimens, 140 spores, Fig. 7a), amygdaloid-fusoid, sometimes with a low suprahilar depression, moderately to fairly strongly verrucose, warts mostly not anastomosing, weakly dextrinoid, yellowish brown. Although narrow, the spores are quite different in appearance than in C. balteaticlavatus. Basidia 30–39 × 7–8.5 µm (50 basidia), clavate, pale brownish, very finely granulose at the base. Lamellar trama hyphae with moderate to large, dark rice-like granules on yellowish green ground colour. Stipe apex hyphae pale yellow to pale brown, outermost hyphae with reddish brown, granulose contents. Pileipellis: epicutis very weakly gelatinous, uppermost hyphae 3–8 µm wide, pale ochraceous to reddish brown, very finely, densely incrusted, with some small, red brown granules. Deep in the pileipellis some hyphae with fairly large, red brown to blackish brown granules and chips. The thin-walled upper hyphae lost with age and are mostly not seen in older basidiomes. Hypoderm absent. Ecology & Distribution — Presumably with Betula, in subalpine birch forests but also in middle and northern boreal forests at least in oceanic areas. Produces fruitbodies from mid-August to mid-September. Known from Fennoscandia. No sequences of this species exist in public databases. Other specimens examined. Finland, Inarin Lappi, Utsjoki, Kevo, shore of the river Tsarsjoki, W of the cascade, herb-rich, temporarily flooded site on a small island, with Betula and Salix bushes, 16 Aug. 1995, J. Vauras 10452 (TUR5282). – Norway, Sogn og Fjordane, Sogndal, on the E side of the main road vis-vis the Kaupanger centre, deciduous forest (Populus, Betula) with some Pinus, 10 Sept. 2000, J. Ruotsalainen 100900 (H); Sør-Trøndelag, Oppdal, Ålbu, Ålbusbakkan, sloping dryish forest with Pinus, Corylus and Betula, 5 Sept. 2010, I. Kytövuori (H); Nordland, Grane, Fagerli, rich spruce forest with few young Betula, calcareous soil, 3 Sept. 2010, I. Kytövuori (H). Notes — Cortinarius balteatialutaceus belongs to /Balteatoalbi together with C. balteatoalbus Rob. Henry and C. balteati­ clavatus Kytöv., Liimat. & Niskanen. In the ITS regions it differs by 5 substitutions and indel positions from C. balteatoalbus and 7 substitutions and indel positions from C. balteaticlavatus. The sequences of C. balteatialutaceus have one base difference and the maximum pairwise distance is one. Typical for the species of the clade, compared to many other species of /Phlegmacioides, are basidiomata totally without bluish tints even when young and narrow, rather small spores. Cortinarius balteatialutaceus is a stout, C. balteatus-like species and grows in subalpine birch forests but also in middle and northern boreal forests at Persoonia – Volume 33, 2014 least in oceanic areas. The sister species C. balteatoalbus has less verrucose spores and lamellar trama hyphae are densely small-granulose in MLZ. In addition, C. balteatoalbus has a more southern, hemiboreal to montane distribution although it also rarely occurs in the middle boreal zone. Cortinarius balte­ aticlavatus has a relatively longer stipe, brown pileus margin, on average smaller spores (av. = 8.5–9.3 × 4.9–5.1 µm) and it grows in mixed forests in boreal zone. Cortinarius balteatus (Fr.) Fr. and C. badiolatus (M.M. Moser) M.M. Moser occur in subalpine forests but have larger spores, C. balteatus 10–12 × 5.5–6.5 µm and C. badiolatus 10.5–12.5 × 5.5–7.5 µm. In addition, the pileus margin of C. balteatus is usually blue, at least when young. Cortinarius areni­silvae (Brandrud) Brandrud is reminiscent of C. balteatialutaceus but the former grows in dry, sandy pine heaths and has smaller spores (7.7–9.5 × 4.3–5.2 µm). Cortinarius balteatibulbosus Kytöv., Niskanen, Liimat., Bojantchev & A.F.S. Taylor, sp. nov. — MycoBank MB805878; Fig. 6e, 7b Etymology. The name refers to the confusion of this species with C. bal­ teatoalbus and to the marginate bulb. Type. Finland, Uusimaa, Espoo, Nuuksio, Pirttimäki, half-open cut meadow, 4 Sept. 1998, I. Kytövuori 98-1624, H6033539 (holotype H; isotype NY). GenBank KF732589. Pileus 6–13 cm broad, hemispherical to plano-convex, slightly viscid in moist weather, otherwise dry, coarsely innately fibrillose, pale brown with a whitish to very pale brown margin, rarely completely white, becoming brown with age. Lamellae emarginate, crowded, pale grey when young, later pale brown. Stipe 4–7 cm long, 1.5–2.5 cm thick at apex, 2.5–4 cm wide at base, with a more or less marginate bulb, whitish. Universal veil white, later ochraceous brown at bulb margin, sparse. Context white. Odour indistinct in lamellae, in flesh mild and pleasant. KOH­reaction in context yellow. Exsiccata: pileus brown to brownish to leathercoloured, stipe at the top whitish, at the base concolorous with the pileus. In MLZ: Spores 9.1–9.9 –10.9 × 5.2–5.7–6.1 µm, av. = 9.6– 10.2 × 5.6 – 5.8 µm, Q = 1.58 –1.74 –1.89, Qav. = 1.66 –1.82 (6 specimens, 120 spores, Fig. 7b), broadly amygdaloid to citriform, fairly strongly, darkly verrucose, fairly dark-coloured, somewhat dextrinoid. Basidia 30–42 × 7–10 µm (60 basidia), clavate, brownish, finely granulose and with few larger granules at the base. Lamellar trama hyphae with moderate to large, dark rice-like granules on brownish to yellowish ground colour. Stipe apex hyphae orange brown, in outermost hyphae with abundant dark red brown, partly blackish brown contents. Pileipellis: gelatinous layer practically absent, uppermost hyphae 4–8 µm wide, thin-walled, ochraceous brown to reddish brown, mostly finely, densely, spirally incrusted, with few red brown granules. Lower hyphae somewhat wider with slightly thicker, smooth to incrusted, brown walls and dark sepia brown granular contents, granules small to large. The thin-walled upper hyphae lost with age and are mostly not seen in older basidiomata. Ecology & Distribution — In rich mixed forests, in parks, and on yard lawns with different deciduous trees (Tilia, Quercus, Fagus, Populus, Corylus, Betula). Until now all the collections known to us are from the hemiboreal zone. Widespread in Europe, from Bulgaria to Finland. Can fruit very early in the season, in July, but also in September. Other specimens examined. Bulgaria, Saranzi, 700 m asl, Quercus cerris, Q. petraea and Q. frainetto, 10 June 2010, D. Bojantchev DBB33060 (UC). – Finland, Varsinais-Suomi, Vihti, Lintumäki, Ruohoisnummentie, fairly old, fairly rich Picea abies dominated forest, 19 July 2004, H. Tuovila, H6027358 (H); Uusimaa, Espoo, Nuuksio, Pirttimäki fairly rich grass-herb forest with Populus tremula, Betula, Alnus incana, Quercus, Corylus, Prunus padus, Salix spp., and some old pines and young spruces, 2 Sept. 1993, I. Kytövuori 93-639, K. Liimatainen et al.: Identification and nomenclature of Phlegmacium H6032755 (H), 4 Sept. 2004, Ukkola (H); Helsinki, Tamminiemi, on yard, under Tilia, on lawn, 3 July 2000, T. Niskanen H6032413 (H); cut lawn, under Tilia, 18 Sept. 2000, I. Kytövuori (H); Vantaa, Tikkurila, at the river, on a cut lawn under Tilia, 22 July 2004, I. Kytövuori 04-044 (H). – Sweden, Uppsala, Berthåga graveyard, mixed forest, 21 July 2004, A. Taylor 2004091, UNITE No. UDB001132 as ‘C. balteatoalbus’ (UPS); A. Taylor 2004127, UNITE No. UDB000722 as ‘C. balteatoalbus’ (UPS); Järfälla, Järfälla Naturreservat, mixed forest of Picea abies, Quercus robur, Betula, Corylus, 7 Sept. 2010, D. Bojantchev DBB36892 (UC); Slottsbacken, under Fagus sylvatica, 2 July 2004, A. Taylor 2004045, UNITE No. UDB000711 as ‘C. balteatoalbus’ (UPS); Stenhagen, mixed forest, 10 July 2004, A. Taylor 2004088, UNITE No. UDB000715 as ‘C. balteatoalbus’ (UPS). Notes — Typical for C. balteatibulbosus are low and stout basidiomata without bluish tints, coarsely innately fibrillose pileus, marginate bulb, broadly amygdaloid to citriform, fairly strongly verrucose spores, and habitat with deciduous trees. The species of /Balteatoalbi have smooth to only finely fibrillose pileus and clavate to almost cylindrical stipe. In addition, the spores of C. balteaticlavatus Kytöv., Niskanen & Liimat. are smaller, especially shorter. Cortinarius balteatialutaceus is almost whitish and has narrower spores and a much more northern distribution. Cortinarius balteatoalbus is paler, the spores are narrower, pileipellis is more incrusted, and in MLZ the red brown small granules (large, dark ones in C. balteatibulbosus) are abundant in lamellar trama hyphae. In our phylogenetic analysis C. balteatibulbosus forms a well supported clade in /Phlegmacioides. It groups together with C. flavescentipes Reumaux (= C. balteatocumatilis Rob. Henry ex P.D. Orton s.auct.) and C. pseudonebularis Moënne-Locc. (1.00 PP). The sequences of C. balteatibulbosus have one length polymorphism and the maximum pairwise distance is zero. In ITS regions it differs by 19 substitutions and indel positions from C. flavescentipes. The species was previously called C. balteatoalbus in northern Europe but our studies revealed that C. balteatoalbus is a completely different species and therefore this one is described as new. Cortinarius balteaticlavatus Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805879; Fig. 6f, 7c Etymology. The name refers to the affinity of C. balteatus and clavate stipe. Type. Finland, Pohjois-Häme, Virrat, Hauhuu, Sikosaari, Salmela, on the grassy roadside with young Picea, Betula, Populus tremula, Alnus incana and Salix spp., 23 Aug. 1996, I. Kytövuori 96-595, H6032412 (holotype H; isotype NY). GenBank KF732596. Pileus 5–9 cm broad, hemispherical to plano-convex, with an involute margin when young, dry, sand brown to brown, com- a 135 pletely without bluish tints. Lamellae emarginate, crowded, pale brownish grey when young, later brown. Stipe 6–11 cm long, 1.2–2 cm thick at apex, 1.5–2.5 cm wide at base, clavate, whitish. Universal veil whitish to ochraceous, fairly sparse. Context thick, white to very pale marbled brownish. Odour indistinct. KOH­reaction in context yellow. Exsiccata: pileus uniformly pale brown to brown, stipe whitish at the top, pale brown at the base. In MLZ: Spores 8.2–8.9–10.0 × 4.5–5.0–5.4 µm, av. = 8.5–9.3 × 4.9–5.1 µm, Q = 1.60–1.78–1.94, Qav. = 1.72–1.85 (5 specimens, 160 spores, Fig. 7c), narrowly amygdaloid to fusoid, with blunt apex, finely, densely verrucose, warts not anastomosing, fairly light-coloured, very faintly dextrinoid. Basidia 27–39 × 6.5–8.5 µm (40 basidia), clavate, 4-spored, pale brownish, finely granulose at the base, sometimes with a few dark granules. Lamellar trama hyphae with small to moderate, dark rice-shaped granules. Stipe apex hyphae yellowish brownish, outermost ones with yellow brown to red brown substance and some dark granules. Pileipellis: gelatinous layer absent, uppermost hyphae 4–10 µm wide, thin-walled, ochraceous brown, finely, densely, spirally to spot-like incrusted, with many large, dark red brown, angular particles. Lower hyphae with slightly thicker, smooth to incrusted, brown walls and red brown to sepia brown granular contents, granules small to large. The thin-walled upper hyphae lost with age and are mostly not seen in older basidiomata. Ecology & Distribution — In mixed forests (Betula, Populus, Salix, Picea, Pinus) but the tree host is unknown, most of the collections from grassy roadsides. Occurs in the southern boreal to the northern boreal zones, known from South Finland to Lapland. Fruits from mid-August to mid-September. Other specimens examined. Finland, Etelä-Savo, Mäntyharju, Juolasvesi, mesic spruce heath forest with swampy depressions, Pinus, Betula and Populus tremula, road ditch, 2 Sept. 1996, I. Kytövuori 96-782, H6032415 (H); Salmela, half-open forest of Betula and Populus tremula and some pines and spruces, 22 Aug. 1998, I. Kytövuori 98-1337 (H); Koillismaa, Taivalkoski Jurmu-Kurtti, Koivuoja, pine dominated forest with some Picea and Betula, by a forest track, 31 Aug. 2008, I. Kytövuori 08-584, H6033533 (H); Inarin Lappi, Inari, N side of the river Lutto, N side of the road to Rajajooseppi, Keskikompsio (Hätäpuhelin), above the river bank, mesic heath forest with swamp depressions, Pinus, Betula and Salix spp., 14 Aug. 1995, I. Kytövuori 95-382, H6032729 (H). Notes — Cortinarius balteaticlavatus belongs to /Balteatoalbi together with C. balteatoalbus Rob. Henry and C. balte­ atialutaceus. In ITS regions it differs by 10 substitutions and indel positions from C. balteatoalbus and 7 substitutions and indel positions from C. balteatialutaceus. The sequences of C. balteaticlavatus are identical. Typical for the species of the c b d f e Fig. 7 Spores of: a. C. balteatialutaceus IK09-751; b. C. balteatibulbosus IK98-1624; c. C. balteaticlavatus IK96-595; d. C. brunneiaurantius JV17979; e. C. caesiocolor IK00-029; f. C. myrtilliphilus IK 97-1469, in Melzer’s reagent. — Drawn by I. Kytövuori and T. Niskanen. — Scale bar = 10 µm. 136 clade, compared to many other species of /Phlegmacioides, are basidiomata without bluish tints and narrow, rather small spores. Cortinarius balteaticlavatus grows in mixed forests in boreal zones. It is a stout species that resembles C. crassus Fr. The latter, however, has narrower spores (7–9 × 3.5–4.5 µm), abundant cylindrical to clavate cystidia especially at the lamellar edge, and no KOH­reaction in context. Cortinarius balteatoalbus differs from C. balteaticlavatus by its lower and wider appearance, larger spores, and more incrusted pileipellis hyphae. Cortinarius balteatialutaceus has a relatively shorter stipe, whitish to very pale brown pileus margin, and larger, moderately to fairly strongly verrucose spores. Cortinarius brunneiaurantius Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805880; Fig. 6g, 7d Etymology. The name refers to the colour of the pileus. Type. Finland, Varsinais-Suomi, Turku, Ruissalo, Kansanpuisto, Tilia alley, also Quercus robur nearby, on clayey-mull soil, 22 Sept. 2001, J. Vauras 17979 (holotype H6032422; isotype TUR171972). GenBank KF732600. Pileus 4–9 cm broad, hemispherical to plano-convex, at first viscid but soon dry, centre with small, appressed scales, innately fibrillose toward margin, pale ochraceous brown. Lamellae emarginate, crowded, bluish to pale grey when young, later pale brown. Stipe 4–9 cm long, 0.8–1.5 cm thick at apex, 1.2–2 cm wide at base, clavate, first whitish, later pale brown. Universal veil white, soon brownish, rather sparse. Context white in the pileus, bluish at least in the upper part of the stipe. Odour not recorded. KOH­reaction in context yellow. Exsiccata: pileus orange brown to brown at the centre, paler yellowish brown at the margin, stipe pale brown. In MLZ: Spores 8.2– 8.9 –9.7 × 4.8–5.2 –5.4 µm, av. = 8.9 × 5.2 µm, Q = 1.58–1.71–1.83, Qav. = 1.70–1.71 (2 specimens, 100 spores, Fig. 7d), amygdaloid to citriform, with a shallow suprahilar depression, moderately to fairly strongly verrusose, warts dark, moderately dextrinoid, often very dark. Basidia 30–38 × 7–8 µm, 4-spored, clavate, sand brown, with very small granules. Lamellar trama hyphae with dark small granules and blood red substance or large granules and chips on greenish greyish ground colour. Stipe apex hyphae bright yellow, some with granules and chips, outermost ones yellow to reddish brown with blood blackish contents. Pileipellis simplex, in overall view orange red, uppermost hyphae 4–8 µm wide, with clearly discernible spirally incrusted wall and some dark granule mounds inside. Lower hyphae wider, more yellow, almost smooth, with abundant dark granular substance. Hypoderm absent. Ecology & Distribution — In deciduous forests, presumably with Quercus, Corylus and Populus. To date only known from South Finland. Fruits in autumn. No sequences of the species exist in the public databases. Other specimens examined. Finland, Uusimaa, Espoo, Nuuksio, Pirttimäki, fairly rich grass-herb forest with Populus tremula, Betula, Alnus inca­ na, Quercus, Corylus, Prunus padus, Salix spp., and some old pines and young spruces, 2 Sept. 1993, I. Kytövuori 93-644, H6032406 (H). Notes — Cortinarius brunneiaurantius belongs to /Phlegmacioides and forms a well supported group (0.99 PP) with C. sobrius P. Karst., C. balteatus (Fr.) Fr., C. brunneoviolaceus Bidaud, C. pseudonaeovosus Rob. Henry and C. clarobaltoides var. longispermus Reumaux. The ITS sequences of C. brun­ neiaurantius are identical and differ from the ones of the sister species C. sobrius by 9 substitutions and indel positions. Morphologically, it does not resemble any of its closest relatives. Typical for the species are pale ochraceous brown pileus, rather small, amygdaloid spores, positive granular MLZ reaction, and habitat with deciduous trees. Persoonia – Volume 33, 2014 Cortinarius caesiocolor Kytöv., Liimat. & Niskanen, sp. nov. — MycoBank MB805881; Fig. 6h, 7e Etymology. The name refers to the colour of the pileus. Type. Finland, Uusimaa, Lohja, Jalassaari, Tamminiemi, by a track with Betula, Populus tremula, Quercus, Corylus and Salix caprea, on calcareous soil, 27 Aug. 2000, I. Kytövuori 00-029 (holotype H; isotype NY). GenBank KF732603. Pileus 5–9 cm broad, hemispherical to convex, then planoconvex, slightly viscid, innately fibrillose, more so toward margin, bluish violet to violet brown. Lamellae emarginate, crowded, first pale greyish brown with a bluish tint, later pale brown to brown. Stipe 6–10 cm long, 1–2 cm thick at apex, 2–3.5 cm wide at base, clavate, whitish, with a bluish tint at the apex. Universal veil first pale blue, later becoming brown, sparse. Context blue in all parts or partially whitish but blue at least close to lamellae. Odour indistinct. KOH­reaction in context yellow. Exsiccata: pileus pale greyish brown to violet brown, stipe pale greyish brown, brown at base. In MLZ: Spores 9.1–9.9–10.9 × 5.4–5.8–6.3 µm, av. = 9.8–9.9 × 5.7– 5.9 µm, Q = 1.59 –1.70 –1.84, Qav. = 1.67–1.72 (2 specimens, 120 spores, Fig. 7e), weakly citriform, sometimes with a low suprahilar depression, dark-coloured, moderately dextrinoid, fairly strongly verrucose, warts anastomosing, dark. Basidia 30–41 × 7.5–9.0 µm (40 basidia), 4-spored, clavate, reddish sand brown, mostly with some blood red particles. Lamellar trama hyphae yellow to orange yellowish, with small granules. Stipe apex hyphae yellow to orange yellow, with small granules, the outermost ones fairly red, with very small blood red granules. Velum/cortina hyphae fairly red, with very small blood red granules. Pileipellis: epicutis weakly gelatinous, uppermost hyphae narrow, 2–4 µm wide, ochraceous brown, spirally incrusted, mostly not granulose, mostly without large redbrown particles, lower hyphae up to 8 µm wide, mostly incrusted, somewhat granulose with small, brown granules. Hypoderm absent. Ecology & Distribution — Under Quercus, Populus and Co­ rylus on mull soil in parks, roadsides and deciduous forests. Known from southern Finland. No sequences of this species exist in public databases. Other specimen examined. Finland, Uusimaa, Helsinki, the cemetery of Malmi, Quercus robur alley, on cut meadow, 17 Aug. 1997, I. Kytövuori 97207, H6032730 (H). Notes — Cortinarius caesiocolor is a typical member of /Phlegmacioides. Typical for the species are bluish to violet pileus and upper part of the stipe (appearance somewhat like C. porphyropus), small spores, and habitat with deciduous trees. The sister species C. chromataphilus Rob. Henry and more distantly related C. largus Fr., which also grow in deciduous forests, are at pileus centre greyish brown, towards margin greyish blue, and have larger spores (spores of C. chromataphilus 10.5–13 × 6–6.5 µm, of C. largus 10.0–11.8 × 6.1–8.8 µm). Another small-spored species with bluish colours is C. violaceomaculatus Brandrud, but it has smaller (8.8–10.2 × 5.2–5.7 µm), more weakly verrucose spores, and grows with conifers. Cortinarius caesiocolor formed a well-supported clade in our phylogenetic analysis and differs in ITS regions from C. chromataphilus by 16 substitutions and indel positions. The sequences of C. caesiocolor have three base and one length polymorphisms and the maximum pairwise distance is zero. Cortinarius myrtilliphilus Kytöv., Liimat., Niskanen & Brandrud, sp. nov. — MycoBank MB805882; Fig. 7f Etymology. The name refers to the habitat with Vaccinium myrtillus. = Cortinarius vacciniophilus Brandrud, Edinburgh J. Bot. 54, 1: 114. 1997. p.p. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium Type. Finland, Kainuu, Suomussalmi, Suolijärvi, Siikajärvi, NW foot of Siikavaara (Rönninvaara), Pöksäkorpi, gently sloping, partly swampy, grassherb-spruce forest with some Pinus, Betula, Populus, Alnus incana and Salix spp., 14 Sept. 1997, I. Kytövuori 97-1469, H6032751 (holotype H; isotype NY). GenBank KF732605. Pileus 4.5–9(–11) cm broad, hemispherical to plano-convex, with slightly incurved margin, weakly to distinctly viscid when young, soon dry, with appressed veil scales, especially at centre, pale yellowish brown to darker ochraceous brown to leather brown (like C. balteatus), young margin paler, almost whitish; sometimes with hygrophanous streaks. Lamellae emarginate, crowded, greyish white when young, later pale brown. Stipe 5.5–11 cm long, 1–2(–2.5) cm thick at apex, 1.5–3 cm wide at base, clavate, at first white, later becoming pale brown from base. Universal veil white, sparse. Context white, faintly greyish hygrophanous at stipe apex. Odour indistinct to weak, pleasant, resembling corn. KOH­reaction in context strongly yellow when young, later weakly yellow or with yellow margin. Exsiccata: pileus uniformly pale warm yellowish brown to brown or somewhat darker at the middle, stipe with the same tint as pileus but lighter. In MLZ: Spores 10.9–12.1–13.4 × 6.3–6.9 –7.5 µm, Q = 1.63– 1.79 –1.91 (2 specimens, 100 spores, Fig. 7f), amygdaloid-fusoid with a distinct suprahilar depression and a long apical part (in form very much like those in C. collinitus), (finely to) moderately verrucose, somewhat dextrinoid. Basidia 33–45 × 8.5–11 µm (40 basidia), sand brown, dark granules very few. Lamellar trama hyphae with small granules. Stipe apex hyphae: pale brown, outermost hyphae more reddish, dark granules few. Pileipellis: gelatinous layer present, erect-sinuose gelatinous hyphae 4–6(–8) µm wide, repent subsurface hyphae 5–10 µm wide, fairly thin-walled, finely to fairly strongly spirally incrusted, with some dark red brown, angular particles. Lower hyphae with slightly thicker, smooth to strongly and coarsely incrusted hyphae with dark red brown, farinose to granulose contents. Ecology & Distribution — In mesic Picea dominated forests, from richer to poor Vaccinium myrtillus type. So far only known from middle boreal zone from Finland and Norway and might be rare. Produces fruitbodies in autumn. Other specimens examined. Norway, Oppland, Ringebu, Venabygd, c. 700 m asl, Picea abies forest, ± poor raw humus, mossy Vaccinium myrtillus type, 30 Aug. 1994, E. Bendiksen & T.E. Brandrud, TEB 4-94, O 125949 (O, sub nom. C. vacciniophilus). Notes — Cortinarius myrtilliphilus belongs to /Phlegmacioides. It has an ochraceous brown pileus and pale greyish white lamellae, both without bluish tints, large spores, and it grows in mesic Picea dominated forests. It has previously been confused with C. pseudonaevosus Rob. Henry (= C. acidophilus Brandrud) which is a common spruce forest species in northern Europe and also occurs in mountain areas of Central Europe. Typically, C. pseudonaevosus has bluish pileus margin, bluish tints in context of the pileus and stipe apex, and smaller spores. However, C. myrtilliphilus strongly resembles the large-spored and non-bluish specimens of C. pseudonaevosus. Previously, these variants were called C. vacciniophilus Brandrud (Brandrud 1998). These large-spored C. vacciniophilus specimens are in some regions as frequent as the smaller spored C. aci­ dophilus specimens (Gjerde et al. 2012). Our molecular studies showed that a majority of the large-spored specimens are in fact C. pseudonaevosus, leaving C. pseudonaevosus as an apparently genetic uniform but morphologically heterogeneous species. On the other hand, analysis of ITS sequences also showed that C. myrtilliphilus represents an autonomous species and C. myrtilliphilus and C. pseudonaevosus are not closely related. Cortinarius pseudonaevosus belongs to a strongly supported clade (0.99 PP) with C. balteatus (Fr.) Fr., C. brunneiaurantius Kytöv., Liimat. & Niskanen, C. brunneoviolaceus Bidaud, C. so­ brius and C. clarobaltoides var. longispermus Reumaux, while 137 C. myrtilliphilus is placed elsewhere in /Phlegmacioides, but further relationships with the other members of the clade were not fully resolved. Based on the pairwise comparison of the ITS sequences the closest species is C. badiolatus (M.M. Moser) M.M. Moser (= C. durus s.auct.) from which it differs by 11 substitutions and indel positions. Cortinarius badiolatus differs from C. myrtilliphilus by paler, initially almost whitish colours of the pileus, and habitat mainly in subalpine Betula forests. Based on molecular and morphological data we conclude that C. myrtilliphilus represent an autonomous species and describe it here as new to science. Cortinarius myrtilliphilus must be a very rare species, by us only found twice during many years of study. Due to limited material, we do not fully know the morphological variation of C. myrtilliphilus, and the morphological overlap with the similar, but genetically quite distant, largespored variants of C. pseudonaevosus (= C. vacciniophilus p.p.) needs further study. DISCUSSION Type studies The majority of names published in the genus Cortinarius have been by French taxonomists, representing over half of the types we studied, 132 names representing 77 species, but for 17 of these an earlier name by other taxonomists exist, so in reality the number of truly new species they described is 60. They represent species from both broadleaf hardwood (about 60 %) and conifer forests. Thus, the Cortinarii of France are among the most extensively studied in the world. Although Friesian names often dominate the nomenclatural discussions in Cortinarius, the number of species he described is significantly less than the numbers for French authors. In Fries (1836 –1838) there are about 50 names belonging to the groups we studied and of these 32 are currently included in Funga Nordica (Jeppesen et al. 2012). This demonstrates how difficult it is to interpret old names with vague descriptions and no type material. The Friesian names currently in use include mainly species for which either published or unpublished illustrations exists making them easier to interpret. Species with Friesian names represent about 20 % of the species recognized in this study. A similar trend occurs in subg. Telamonia. For example, in sections Armillati and Brunnei 30 % of the total known species were described by Fries, and in the morphologically challenging sect. Bovini only 8 % (Niskanen et al. 2009, 2011a, 2013a). From North America we studied 60 types, representing 59 species. Of these, 52 species (88 %) have apparently not been described from anywhere else in the world, while one of them is the older synonym for a European name (C. metarius Kauffman = C. barbarorum Bidaud, Moënne-Locc. & Reumaux). Thirty-five of these species were described from the mountains and coastal areas of western North America, 13 species are from the Rocky Mountains and 12 species were collected from eastern North America, mainly Michigan. Almost all the names that are synonyms of European species are associated with conifers, i.e., C. crassus Fr. (= C. subaustralis A.H. Sm. & Hesler, described from North Carolina), C. glaucopus (Schaeff.) Fr. (= C. glaucopoides Kauffman, described from Colorado) and C. metarius; the only exception is C. triumphans Fr. s.auct. (= C. ophiopus Peck from Maryland). Species described from North America and reported here for the first time from Europe, include C. olympianus (USA, Washington state and Finland), C. patrickensis (USA, California and Sweden) and C. sannio (USA, Wyoming, Finland and Sweden), all are conifer-associated species. These findings correlate well with previous studies by Garnica et al. (2011), Harrower et al. (2011) and Niskanen et al. (2013a, b). Interestingly, many names described from Europe 138 have been used in North America, whereas few names from North America have been applied to taxa in Europe. Of the 154 species recognised in this study 114 of them have no synonyms, however, the remaining 40 species each have one or more synonyms. The species with a significant number of synonyms include C. largus (14), C. pseudonaevosus (6), C. talus (5), C. crassus (4) and C. varius (4), all represent rather common species. The authors whose names we studied have all described synonyms. This is in part due to the difficulty in determining which species already have been described, especially based only on morphology and ecology. The narrow species concept of Bidaud, Henry, Moënne-Loccoz and Remaux was not supported by our study nor were the broad species concepts of Fries, Brandrud and Jeppesen et al. (2012) as supported by the results of Frøslev et al. (2007) and Niskanen et al. (2009, 2011a). In this paper, species names have been synonymised when both molecular and morphological data have supported it. Of course there is some risk that morphological differences have been overlooked by studying too few specimens. Based on the overall data we have on Cortinarius it would seem logical, however, that in the majority of cases the synonymization is correct. In doubtful cases, when the variation in ITS regions of a group of specimens has been more than 2 substitutions and/ or indel positions and we have not had enough specimens to make further comparisons, we have left names unchanged, i.e. C. albescens/C. gentianeus/C. volvatus, C. herpeticus/C. vio­ laceonitens, and C. misermontii /C. olidoamarus var. valentinus / C. subaccedens/C. van­campiae. Also, the following species with higher intraspecific variation may in reality represent two to several species: C. aureofulvus s.auct. /C. orichalceus var. xan­ thocephalus, C. pallidirimosus, C. porphyropus and C. scaurus. Our study includes the majority of names described in subg. Phlegmacium, excluding sect. Riederi and parts of sections Calochroi and Fulvi already treated or to be treated by other taxonomists (e.g. Frøslev et al. 2007). In some instances, type material could not be acquired or found in herbaria, i.e. the material of C.H. Peck (NYSM) and some of the material of R. Henry (PC). In other cases they were not possible to study within the time frame of this study, i.e. the early names described by M. Moser (M). Peck published from 1870 to 1912 about 80 names in the genus Cortinarius and the names represent at least twenty species of subg. Phlegmacium s.lato based on morphological and microscopical features. The material deposited in München includes about 30 Phlegmacium species. For about 55 Phlegmacium species described by Henry the type material was not found and over 20 additional ones remain to be studied. Therefore, it is possible that some of these names have priority over currently applied Cortinarius names, including some of those in this paper. Our plan is to acquire Peck’s, the remaining materials of Moser’s and Henry’s, and perhaps others that we have overlooked in the near future and attempt to sequence them for comparison. New species The majority of new species described here represent boreal taxa, i.e. C. pallidirimosus, C. balteaticlavatus and C. myrtil­ liphilus, and/or are species that previously have been included within the morphological limits of other, ‘well-known’ taxa, these include C. boreicyanites and C. subfraudulosus. The large number of unknown, boreal species is not surprising, since the majority of existing names are described from hemiboreal and temperate zones or from the mountains of Central and southern Europe. Even though the Cortinarius species of Europe have been extensively studied, there undoubtedly remain a lot of taxa to be discovered, not to mention in other areas of the world, which are less explored. Persoonia – Volume 33, 2014 Factors affecting on the success of molecular work The success of the molecular studies was significantly influenced by the condition of type specimens, including the age of the specimen, how it was dried and/or preserved, and whether or not it was mouldy. Also, the length polymorphism of the ITS region caused some problems with direct sequencing. In general, Cortinarius is one of the easiest genera of Agaricales for molecular work. Often DNA can be extracted with regular methods even from old specimens. For example, we obtained a complete ITS region from the type material fo C. sobrius P. Karst. (collected in 1890) and C. virentophyllus Kauffman (collected in 1912). The success rate usually decreases with older specimens, however, there is some difference between the specimens collected by different authors. For example, we were not able to successfully sequence any of the specimens collected by T. Hongo during 1960 –1968 (Niskanen et al. 2011a, and unpublished data of C. cinnamomeoides Hongo, C. neoarmillatus Hongo and C. nigrosquamosus Hongo) whereas the success rate for the specimens collected by A.H. Smith mainly during 1935 –1941 was much better. This could be due to the drying process itself, for example, drying temperature, length of drying time, which is related to air circulation, and the condition of fresh specimens when placed on the dryer. In particular, the type collections of R. Henry proved challenging, since often the material was sparse and poorly labelled, and many were mouldy. In some instances types were missing, could not be located at the time they were requested, or were not available for study. Several things can improve the success rate of problematic specimens. For example, methods that provided high quality DNA extraction, and PCR and sequencing machines improved the results. If the DNA is fragmented, amplifying and sequencing ITS1 and ITS2 regions separately was often helpful. Recommendations for stabilizing the nomenclature For achieving a stable and unambiguous use of names, the following requirements should be fulfilled whenever possible. Types of existing names should be studied. Even easily identifiable species like C. balteatoalbus can be misinterpreted, or a species may turn out to be a group of cryptic species, for example, C. dionysae and C. elegantior (Garnica et al. 2011). Taxonomists should not use names in barcoding databases unless they have been verified by molecular type studies. For old names that do not have type specimens or where the type specimen is in poor condition or not available because of historical preservation, a neotype, lectotype or epitype should be chosen. Names rejected due to the problems of interpretation should be excluded from consideration. If the name is not verified with a sequence from a type specimen its application will be uncertain and a source of confusion. A significant problem with neo-, lecto- and epitypes is the lack of a database for them. It is difficult to find out if a type already has been designated thus hindering nomenclatorial work. New names should not be accepted for publication without the support of molecular data in genera where the amplification of the ITS or other sequences is easily done, thus preventing the description of synonyms and creating a sound reference database for further taxonomical and ecological studies. Finally, when studying types it is essential to mark the basidiomata from which the DNA has been extracted. Some collection might turn out to be mixed, and thus, knowing exactly which basidiocarp has been studied, will be necessary. These recommendations may seem self-evident, but have been neglected in many cases. Often the reason is a lack of resources and/or knowledge. At least in Cortinarius the amount of already existing names is overwhelming and we need to proceed carefully from this point forward. K. Liimatainen et al.: Identification and nomenclature of Phlegmacium Infrageneric classification The classification of Cortinarius is not yet stabilised and many traditional infrageneric groups have shown to be at least partly artificial. Therefore, when studying a certain group of species, it might be difficult to find all the relevant species described from literature. The aim of the phylogenetic estimates in this study was not to solve the infrageneric classification of Cortinarius but to show the preliminary placement of the studied species, which then could be used for guidance in further taxonomical studies. In our phylogenetic analysis ten of the twelve clades representing phlegmacioid species sensu Garnica et al. (2005) were recovered: /Alluti (= Multiformes), /Amarescentes (= Infracti), /Arguti/Calochroi, /Percomes, /Phlegmacium, /Phlegmacioides, /Praestantes (= Claricolores), and /Scauri. In addition, /Glaucopodes was well supported in our analysis. Clades /Caerulescentes and /Vulpini were split. Barcoding Cortinarius This study provides ITS barcodes for 175 Cortinarius species. This data enables the identification of a majority of boreal and temperate species of Phlegmacia, except for sections Calochroi, Fulvi, for which part of the ITS sequences already are published (e.g. Frøslev et al. 2007) and Riederi, from Europe and parts of North America. Of the 236 names we studied, only 61 were currently represented in GenBank. It is noteworthy that almost half of these names are Friesian names and a third of them are names published from North America. Only 4 % of the names described by French authors have made their way into general use in GenBank. This emphasizes the importance of type studies and the role of taxonomists in the creation of an identification database. If we do not produce sound basic data on Cortinarius species, it is difficult or impossible in many cases for non-taxonomists to identify specimens or environmental samples. A barcoding database based on type studies is essential for ecological, environmental or further taxonomic research. Once completed, it will create a solid base for future studies. Barcoding is a powerful tool, which enables us to identify and compare species from different regions in a completely new way. This is especially true and important for fungi like Cortinarius, which are morphologically challenging and difficult to identify. Finally, we are able to reliably compare our knowledge on species from different areas. A base for Phlegmacia is created here and provides a beginning and direction for future studies in Cortinarius. Acknowledgements We are grateful to the curators of C, G, H, IB, MICH, PC and S, and especially to Bart Buyck, Xavier Carteret, Philippe Clerc, Regina Kühner-Winkler, Ursula Peintner and Patricia Rogers. Anna-Lena Anderberg is also warmly thanked for providing photos of unpublished plates of Fries. Tor Erik Brandrud is thanked for valuable comments concerning the neo- and epitypifications and providing the photo of C. caesiophylloides. Andy Taylor and Dimitar Bojantchev are thanked for providing material and sequences for the study and Andy Taylor also for providing the photo of C. talimultiformis. Ertugrul Sesli is thanked for providing material of C. talimulti­ formis, Tapio Kekki for providing the photo of C. flavipallens, Jukka Vauras for providing the photo of C. brunneiaurantius and Karen Hughes for providing an additional sequence of C. acystidiosus. Bálint Dima and Tobias G. Frøslev thank members of the ‘J.E.C. Phlegmacium Research Group’ (Francesco Bellù, Tor Erik Brandrud, Bernhard Oertel, Günter Saar and Geert SchmidtStohn) and Thomas Stjernegaard Jeppesen, and László Albert for valuable taxonomic discussions during the past years. László Albert and Zoltán Lukács are thanked for making older literature available for study. Finally, we would like to thank the reviewers for valuable comments which helped us to improve the manuscript. 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