Cortinarius Section Bicolores and Section Saturnini (Basidiomycota, Agaricales), a Morphogenetic Overview of European and North American Species
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Persoonia 39, 2017: 175–200 ISSN (Online) 1878-9080 www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE https://doi.org/10.3767/persoonia.2017.39.08 Cortinarius section Bicolores and section Saturnini (Basidiomycota, Agaricales), a morphogenetic overview of European and North American species K. Liimatainen1, X. Carteret 2, B. Dima3,4, I. Kytövuori5, A. Bidaud6, P. Reumaux7, T. Niskanen1, J.F. Ammirati8, J.-M. Bellanger9 Key words Abstract Cortinarius is the largest genus of ectomycorrhizal fungi worldwide. Recent molecular studies have shown high levels of morphological homoplasy within the genus. Importantly, DNA phylogenies can reveal characteristics Bicolores that have been either over- or underemphasized in taxonomic studies. Here we sequenced and phylogenetically Cortinarius phylogeny analysed a large set of pan-European and North American collections taxonomically studied and placed in Cortinarius integrative taxonomy sect. Bicolores and sect. Saturnini, according to traditional morpho-anatomical criteria. Our goal was to circumscribe Saturnini the evolutionary boundaries of the two sections, to stabilize both the limits and nomenclature of relevant species, Telamonia and to identify described taxa which, according to our current understanding, belong to other lineages. Our analysis resolves two clades: /Bicolores, including 12 species, one of which is new to science, and /Saturnini, including 6 species. Fifteen binomials, traditionally treated in these two sections based on morphology, do not belong to the above two phylogenetic clades. Instead, six of these latter are clearly placed in other clades that represent sect. Bovini, sect. Sciophylli, sect. Duracini and sect. Brunneotincti. The presence or absence of blue pigments and the detection of specific odours emerge as clearly misleading taxonomic features, but more surprisingly, spore size and ecology can be misleading as well. A total of 63 type specimens were sequenced, 4 neotypes and 2 epitypes are proposed here, and 1 new combination is made. Article info Received: 13 October 2016; Accepted: 1 May 2017; Published: 10 August 2017. INTRODUCTION French authors (e.g., Liimatainen et al. 2014a). Importantly, by identifying evolutionary units that are independent of morpho- Cortinarius is the largest genus of ectomycorrhizal fungi world- anatomical and ecological traits, DNA phylogenies revealed wide, with no less than 4 701 reported taxa (3 360 species, characters that have been overemphasized in monographic 1 341 infraspecific taxa, http://www.catalogueoflife.org, 28 Sept. studies but also uncovered significant taxonomic information 2016 release). However, the number of species greatly varies that has been neglected by previous investigators (Bellanger et depending on the morphological species concept accepted al. 2015, Loizides et al. 2016). The use of these modern tools a by classical authors. Currently, the two major monographs posteriori, to test the autonomy of previously defined morpho- dedicated to the genus are Cortinarius, Flora Photographica logical species, has been instrumental in delineating objective (CFP), which includes ± 300 species, mostly from northern boundaries to taxa, and when applied to type material, stabi- Europe (Brandrud et al. 2014), and the Atlas des Cortinaires lizes taxonomy and nomenclature at the genus level (Frøslev (ADC), still on-going and which so far recognizes ± 2 500 spe- et al. 2007, Liimatainen et al. 2014b, Cripps et al. 2015). The cies, varieties and forms, mostly from France (Bidaud et al. next challenge of this nascent integrative systematics era is 2015). Recent molecular studies have unveiled high levels of undoubtedly to synchronize the two sources of knowledge, so morphological homoplasy as well as numerous cryptic species that on-going monographs introduce morphogenetic species, within the genus, and as a result, do not support the broad i.e., taxa that are both assigned formal diagnosis and a unique species concept of Scandinavian authors or the narrow one of molecular signature. Historically, mycologists have attempted to tackle the complexity 1 Jodrell Laboratory, Royal Botanic Gardens, Kew, Surrey, TW9 3AB, United of Cortinarius by organizing species in hierarchical infrageneric Kingdom. taxa defined on supposedly stable sets of characteristics 2 68, rue Alexis Maneyrol, F-92370 Chaville, France. (Kühner & Romagnesi 1953, Moser 1967, Melot 1990, Moënne- 3 Department of Plant Anatomy, Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, H-1117 Budapest, Hungary. Loccoz & Reumaux 1990). In spite of their practical application, 4 Department of Biosciences, Plant Biology, P.O. Box 65, FI-00014 University most of these lower level taxonomic divisions have proven to of Helsinki, Finland. be artificial when placed under evolutionary scrutiny (Garnica 5 Botanical Museum, University of Helsinki, Helsinki, P.O. Box 7, FI-00014 et al. 2005). Subgenus Telamonia, however, breaks this rule Finland. as most of the numerous species known to date that produce 6 2436, route de Brailles, F-38510 Vézeronce-Curtin, France. 7 84, avenue de Wagram, F-75017 Paris, France. dry-capped basidiomata lacking vivid colours – the morphologi- 8 Department of Biology, University of Washington, Box 351800, Seattle, WA cal definition of the subgenus and excluding a few sections as 98195-1800, USA. sect. Obtusi, Balaustini, Illumini – form a strongly supported 9 CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry monophyletic clade in all published molecular studies (Peintner Montpellier, EPHE, INSERM, 1919, route de Mende, F-34293 Montpellier Cedex 5, France; et al. 2004, Stensrud et al. 2014). Recently, several sections corresponding author e-mail: [email protected]. within Telamonia have been phylogenetically revised, such as © 2017 Naturalis Biodiversity Center & Westerdijk Fungal Biodiversity Institute 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. 176 Persoonia – Volume 39, 2017 sect. Armillati, Brunnei, Bovini and Disjungendi and more are on ADC (Bidaud et al. 2014, 2015), in which part of the results their way to morphogenetic redefinition (Niskanen et al. 2009, presented here have been incorporated (Table 1). The specific 2011, 2013, Liimatainen et al. 2014a). goals of the present work are: Here we deal with Cortinarius sect. Bicolores and Cortinarius 1. to circumscribe the phylogenetic boundaries of the two sect. Saturnini, which encompass Cortinarius evernius, C. sa- sections, through the analysis of a large internal tran- turninus and their lookalikes. Initially, the two sections were scribed spacer (ITS) rDNA sequence dataset built from distinguished by the extent of veil remnants on the stipe, a pan-European and North American vouchered collections; character considered by some authors to segregate subg. 2. to stabilize the nomenclature and species limits of morpho- Hydrocybe from subg. Telamonia (Moënne-Loccoz & Reumaux genetic Bicolores and Saturnini, through sequencing type 1990). However, this morphological feature may not be support- ed phylogenetically, justifying the revision of the two sections material and designating neotype or epitype when oppor- altogether (Niskanen et al. 2012). Eight to thirty-three species tune; have been described in sect. Bicolores and sect. Saturnini in 3. to assign a molecular signature to the numerous collections the major European monographs, from the pioneering work of taxonomically placed in these two sections in contemporary Kühner & Romagnesi (1953) to the latest two releases of the monographs, but that do not belong in the two clades. Table 1 Cortinarius species classified in sections Bicolores and Saturnini by the main European authors. This study Bidaud et al. (1992, 2014, 2015) Brandrud et al. (1990, 1994, Moser (1967) Kühner & Romagnesi 1998), Niskanen et al. (2012) (1953) Sect. Bicolores Sect. Bicolores Sect. Bicolores Key 3.11.7.6.11 Sect. Bicolores Cortinarius cagei C. minicolor, C. cagei C. bicolor? C. bicolor? C. periodolens ad. int. C. dolabratoides sp. nov. C. dolabratus C. imbutoides C. evernius C. evernius, C. parvulior ad. int. C. evernius C. evernius, C. scutulatus C. evernius C. glaphurus C. tubulosus, C. paranomalus (Sat.) C. hircinosmus C. livor C. livor? C. plumulosus C. fundatus C. bicolor? C. bicolor? C. refectus C. refectus, C. testaceoviolaceus C. bicolor? C. bicolor? C. sp1 C. sp2 C. tortuosus C. tortuosus C. tortuosus C. plumbosus C. tortuosus, C. plumbosus C. turgidipes C. cinnamoviolaceus C. cinnamoviolaceus, C. parevernius, C. imbutus C. cinnamoviolaceus, C. parevernius C. basicyaneus C. parevernius C. disjungendus C. cyanosterix C. mattiae C. mattiae C. mattiae C. subviolascens C. parevernioides C. parevernioides C. salicinus C. salicinus, C. deceptivoides C. quadricolor Sect. Saturnini Sect. Saturnini Sect. Firmiores