sign in sign up
<<
Home , Anamika (fungus), Endoptychum, Gliophorus, Hemileccinum, Hygrocybe procera, Leiotrametes

MYCOLENS (65) , 8 , 14 , Teresa Lebel , Teresa 7 (Nuhn et al. (Nuhn nmark ch. We recommend that that recommend ch. We : [email protected] as a separate for genus as a separate thor e-mail:thor ecvellinga@comcast. (Eberhardt (Eberhardt . 2002), he new taxa proposed, are and this 6 Lille, France , Machiel E. Noordeloos , Machiel , Thomas Læssøe , Thomas 13 6 cal support; (4) different options for the for cal options support; (4) different ylogenetic should evidence be based on e final solutions are properly discussed. properly The are solutions e final However, such phylogenetic analyses phylogenetic such However, molecular-phylogenetic research and the the and research molecular-phylogenetic led has to It trees. phylogenetic resulting cases of characters; many re-evaluation in shown were and gastroid forms sequestrate higher assumed, evolvabilityto have than genera of such subsumption justifying the genera; existing under in for example 1997), Cortinarius & Bruns (Kretzer Suillus et al (Peintner 2015), 2004, Kirk & Verbeken 2007), Lepiota (Ge & (Lebel Tonkin & 2013), and Smith evolvability low had 2013). Lichenization justifying basidiomycetes, recognitionin of Lichenomphalia placed previously three in were species that genera (Redhead et al. 2002). was hoped than extent to a smaller have and delimitation of genus problems solved the several that shown recognition. have Analyses such paraphyletic, genera remain well-known , Alfredo Justo 5 , Gregory M. Mueller 12 et al. 2014, , Roy E. Halling 4 from the Rocky the from Hymenoporus Hymenoporus

commendation how to publish the results of scientific resear of scientific results to publish how the commendation 16 , Pierre-Arthur Moreau , Pierre-Arthur one re 11 et al. 2015). nces pharmaceutiques et biologiques, Université de Lille, 5900 et biologiques, Université nces pharmaceutiques nd information should be included in the publication in which should in t be publication included the in nd information the underlying research is solid, and that the results and th results the is solid, and that research underlying the , Dennis E., Dennis Desjardin The underlying principle for recognition for principle underlying The new The flood these generic recent of 3 ture, phylogenetics, , phylogenetics, ture, n should decision bejustified; final discussed and the the ph (5) from China (Baroni China from paradoxus Tkalčec described based on newly species discovered morphological unique with character are latter of the Examples combinations. crocodilinus Cercopemyces and (USA) Mountains taxonomic rank for that or any of a genus, 1950, 1965). (Hennig is monophyly matter, difficult to extremely was past this the In of fungi, in number as the demonstrate could bemorphological used that characters whether also was unknown limited.was It used, such were that characters some of the or gastroid of sequestrate as formation the justifying evolvability, basidiomes, a low had recognition genera, as separate or a high its downplays which then evolvability, context. a phylogenetic in relevance mainly has been names by prompted nomencla , , Andrew S. Methven , Andrew , and Annemieke Verbeken , and Annemieke , Joe Ammirati , Joe . The 2 10 15 Lyophyllaceae Psathyrellaceae

, was not , was Cortinarius ” for a long time. time. ” for a long , Clark L. Ovrebo, Clark 14 , Thomas W. Kuyper W. , Thomas 1 , P. Brandon Matheny Brandon , P. 9 were proposed. Most of these new of these proposed. Most were basidiomycetes, molecular systematics, molecular basidiomycetes, systematics, , five newin , five genera We formulate five guidelines for introducing new five genera, plus guidelinesintroducing for formulate We

are are for numbers The

Department of Biology, University of Central Oklahoma, Edmond, OK 73034, USA of Central of Biology, University Department of Biology, K.L. Department Ledeganckstraat Belgium 35, 9000 Gent, University, Ghent Département des Sciences Végétales et Fongiques, Faculté des scie Faculté et Fongiques, des Sciences Végétales Département Cook Road, 1000 Lake Chicago IL Glencoe, 60022, USA Garden, Botanic Netherlands 9517, 2300 RA Box Leiden, The P.O. Biodiversity Centre, Naturalis Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, TN USA 37996-1610, Knoxville, of Tennessee, of Ecology Biology,Department University and Evolutionary IL 61920, USA Charleston, of Biological University, Illinois Department Sciences, Eastern Center for Forest Mycology Research, Northern Research Station, USDA-Forest Service, Luquillo, PR USA 00773-1377, USDA-Forest Research Station, Mycology Research, Northern for Forest Center Department of Botany, Institute of Biology, National Autonomous University of Mexico, 04510 Mexico City, Mexico City, 04510 Mexico of Mexico, University Autonomous of Biology, National Institute of Botany, Department 15, 2100 Copenhagen De Ø, of Copenhagen,Universitetsparken University of Denmark, of Biology/Natural History Museum Department 3004, Australia Melbourne Victoria,Royal Gardens Botanic Birdwood Ave, Department of Soil Quality, Wageningen University, P.O. Box 47, 6700 AA Wageningen, The Netherlands; corresponding author e-mail author corresponding Netherlands; The 47, 6700 AAWageningen, Box P.O. University, Wageningen of Soil Quality, Department 98195-1800, USA WA Seattle, of Washington, of Biology, University Department CA USA 94132, San Francisco, Avenue, Holloway 1600 University, State of Biology, San Francisco Department Boulevard, NY Bronx, 10458, USA 2900 Southern Botanical Garden, York New Botany, of Systematic Institute Department of Plant and Microbial Biology, University of California at Berkeley, Berkeley, CA USA; 94720-3102, au corresponding Berkeley, Berkeley, at of California Biology, Microbial and University of Plant Department and three new ones in and three largest far, by genus some 6–7 probably In contrast, affected. under been now subsumed genera have new generic many . With Cortinarius for well-known introduced being names species, it comes as larger an even surprise very few were new genera there to note that generic names encompass species that occur occur species that encompass names generic have and that and Europe America North in been called “Boletus In 2014 and the first six months of six months first 2014 and the In 20 new genera in 2015 alone, than more 25 newcomparable: around counted we same time published that names in generic white-spored,period, of them most with six new genera for species used we to call Clitocybe INTRODUCTION

fungi Six simple guidelines for introducing new genera of genera new for introducing guidelines Six simple net VOLUME 6 · NO. 2 16 12 13 14 15 9 10 11 6 7 8 3 4 5 1 2 Else C. Vellinga Else Jorinde Nuytinck Jorinde Abstract: reviewers and editors adhere to these guidelines. We propose that propose that reviewers and editors guidelines. to these adhere We are: recognized are of species, (1) all geographicsix criteria should genera that be number should tree wide in be coverage phylogenetic monophyletic; (2) the of the coverage,statisti and type sufficient study; species genera to have has under of the phylogenetic trees branching of the (3) the classificatio a formal into tree phylogenetic of the translation one gene; than more and backgrou evidence and (6) all supporting Key words: publication should bepublication peer-reviewed. D. Jean Lodge Jean D. as Boletus (Nuhn et al. 2013, Wu et al., 2014) not. In the case of the boletes, inclusion use these guidelines, but also that reviewers and Psathyrella (e.g, Padamsee et al. 2008). of more sequences from more taxa may and editors of taxonomic journals use these Dealing with the issue of paraphyly then set impact the phylogeny, as the resolution criteria in their assessments of submitted the mycological community on two, rather of the phylogeny of Boletaceae is low manuscripts. divergent, paths. These divergent approaches at many branches (Nuhn et al. 2013, The sixth criterion concerns the way the mirrored old distinctions of splitters and Wu et al. 2014). We strongly advocate results are published and presented to the lumpers. One pathway was to take small that different options are explored and scientific world. MYCOLENS monophyletic groups as the basis for new discussed, instead of using a boilerplate genera, without too much concern about model in which every monophyletic 1. All genera that are recognized the remainder of the original genus. This is clade is translated into a genus. should be monophyletic, not only a practice clearly shown by the treatment of (2) Several of the new genera are erected the one that is the focus of the study, Boletus and Clitocybe. Separation of one small solely based on phylogenetic evidence but also the group from which it is monophyletic group often set into motion a provided by one or two gene regions, separated and the group to which it splitting snowball, as in (Gelardi sometimes only nrITS sequences that is added (the reciprocal monophyly et al. 2015, Vizzini 2015). Unfortunately, do not lend themselves to higher level criterion). recognition of such small genera sometimes phylogenies (Bruns 2001). contributed to the formation of paraphyletic (3) More and more rapid on-line, non- Examples: genera, as in the case of Resupinatus peer-reviewed publications appear (i) When Macrolepiota was split into a which became paraphyletic because of the without any supporting evidence for monophyletic core Macrolepiota with recognition of R. vetlinianus in a separate the newly described taxa. M. procera as its type species, and a genus Lignomyces (Petersen et al. 2015). The second group containing M. rachodes, second approach was taken in Entoloma and We therefore have formulated and present the latter was moved to Clitopilus (Co-David et al. 2009), where here criteria by which the phylogenetic that in itself was only monophyletic by deliberately a broad genus concept was chosen studies and the publications that present also including Endoptychum agaricoides (though other authors opted for smaller genera the data from those studies should be tested (Vellinga et al. 2003) and all necessary (e.g. Largent 1994, Kluting et al. 2014). Other before being accepted. nomenclatorial changes were made examples are Amanita (Justo et al. 2010) (Vellinga 2002. Vellinga & de Kok and Cortinarius (Peintner et al. 2001) where 2002). sequestrate and gastroid species were included PROPOSED GUIDELINES (ii) The genusPorpoloma was found to be in genera with predominantly agaricoid highly polyphyletic and split into four basidiomes. It is an interesting question how Some of our proposed guidelines are genera (Sánchez-Garcia et al. 2014). this divergence should be explained; certainly, so self-explanatory and obvious to us It was essential for the application the existing taxonomy of the group in question that it seems superfluous to present of names to clades to sequence plays a big role. In the boletes, for instance, them here; nevertheless, examples the type species of Porpoloma (see many previously published generic names for that are in conflict with these Recommendation 2). well supported genera, such as and recommendations are surprisingly Strobilomyces, were already available, and the easy to find. The examples we Unfortunately, many studies do not reach proposal of new genera was (consequently) present show how easy it is to meet this standard, as it is very easy to expel considered acceptable. these standards. We realize that species from a genus, without taking into practical issues, such as lost original account the monophyly of the target group. type specimens, or material that Especially in cases of poor resolution PRINCIPLE CONCERNS does not easily yield DNA sequences in a phylogeny, erection of very small may interfere with perfection, monophyletic genera will create more and We find several of the recent trends in but one should at least try, and ever more messy paraphyletic units, from mycological taxonomic research on discuss failures. We emphasize the which the next separation of a new genus basidiomycete fungi disturbing: importance of exploring different can already be predicted. (1) In several groups, the translation from classification options and giving a phylogenetic tree into a classification arguments for the proposed new 2. The coverage of the phylogenetic is taken into extremes, where every taxonomies. The examples we tree has to be broad. single clade is recognized as a separate present are taken from the literature genus. This does not increase insight on basidiomycete fungi, but could Coverage needs to be broad in terms of the: in the evolutionary history of the equally have been chosen from the (a) number of species – it is important to group in question, only inflates the ascomycete literature. remember that a phylogenetic tree only gives taxonomic framework. From a formal The first five criteria relate to the information on those taxa that are included phylogenetic perspective it may not underlying science; the first two criteria in it, and that all statements on relationships matter whether we have one family (e.g. are equally important, and we put less are relative, and not absolute, unless all Boletaceae), with more than a hundred emphasis on the next three. Nevertheless, known taxa are included; (b) geographic genera, or whether we have one it goes without saying that contradiction distribution of the taxa – a phylogeny based genus (e.g. Boletus s. lat.) with many of any of these criteria should be avoided. on species from temperate areas of North infrageneric units, formally named or We recommend first of all that researchers America and Europe, is not informative

(66) IMA MYCOLENS (67) and all full with statistical Albomagister, Pogonoloma, Pogonoloma, , Albomagister, Peer review, though not waterproof, though not waterproof, review, Peer . 2015); and of course, whole genome of course, . 2015); and All supporting evidence and evidence 6. All supporting should be information background which in publication the in included new taxathe are proposed; and should be publication this secondly, peer-reviewed. part first thisThe of promptedguideline is appearanceby the of very short publications officialassociated one of the with taxon supporting any sites, without registration with sometimes evidence nor illustrations, often webpersonal, a link to another, information supporting the site where can be (e.g. found. tree) a phylogenetic is of science results of the Communication of A basic principle event. not a one-time by verifiable are results the is that science go cannot back researchers future in If others. that information, and whole set of data to the be applied. cannot Thisprinciple situation where studies same as experimental in is the used not preserved are cultures the thereby unrepeatable, experiments the rendering are vouchers no reports where field and in past of the publications retained. Paper a source to provide provided and continue be can always consulted; that of information official on line in information that putting library such as depositories JSTORis an extra safe guard. (see debated and always e.g., http://www. nature.com/nature/peerreview/debate/) asis recommended before quality control of reviewpeerpublication. The function is case of taxonomic the novelties in of taxonomic censorship not to exercise to judgedecisions, but new genera against mycological to which the principles these should adhere. community computers, the answer is: taxa answer more the and computers, phylogenies based on genes.more However, only and include rare whole genomes still are of species (e.g. Dentinger number a small et al will becomparisons faced same the with phylogenies as based the issues on a hand-full of genes. Example: nSSU, loci (nLSU, of three an analysis In and rpb2 genes) a wide range from of taxa clade, Sánchez-Garcia tricholomatoid the in et al. (2014) recognized new genera the Corneriella Pseudotricholoma support. Russula and Boletellus Lactarius as a separate as a separate . 2012), and it ; these ; these Heimioporus , beforeRussula , ultimately lat., ultimately s. et al . . (2015) tested different . (2015) tested different . (2014) discussed options . (2014) discussed options . (2008) presented and . (2008) presented et al et al : and Lactarius (2014) discussed the different options options different (2014) discussed the of recognizingfor and consequences one just or or five genus ten genera within opting for a one-genus solution. opting of circumscription for the options of that and delimitation Boletellus regard in genus to form species included that the authors core base of the a grade the at of describing instead genus, that within new genera for each clade. the for options tested 15 different clades different of the phylogeny in up of breaking solution the on settling two genera, into Lactarius , and recognizingand sect. Ochricompactae Multifurca. genus, for recognition of one or three generafor recognition of one or three Porpolomopsis, for Gliophorus, Neohygrocybe The phylogenetic evidence has to has evidence 5. The phylogenetic gene, one be than based more on in genes coding protein preferably SSU- of the regions gene to addition ITS-LSU repeat. are commonly that regionsgene The different all classifications used basidiomycete in and histories, evolutionary different have power resolving different they have hence levels.Phylogenies based only different at not only to be sequences have on nrITS a large with approached dose of skepticism, fact in should no longer bebut accepted basisas for new the genera.marker This barcode for fungal universal speciesis the recognition (Schoch performs this role generally role this performs well because of between large relatedvariation sufficiently not all, but fungal many, species within species-level for that pay groups; we price the over is unalignable nrITS accuracy is that 2001). related taxa (Bruns distantly more better, was past, what debated it was the In taxa/collections more analysed orto have genes taxamore for fewer (Greybeal 1998); and faster costs sequencing lower with classification with monophyletic groups. monophyletic with classification should options be different Therefore and discussed. presented Examples (i) (2014), and Justo & Hibbett Justo (ii) Halling (iii) Buyck (iv) Lodge et al ,

. s. lat.; s.

P. et al Trametes that are always always , hence the the , hence Trametes Chalciporus .

funariophilum P. sect. Marasmius Chalciporus was determined by by determined was et al. 2014). (Jenkinson and occupies an isolated and was shown to was shown Anamika . (2012) refrained from introducing . (2012) refrained introducing from : type all of species genera et al (Nuhn et al. 2013). (Nuhn in the phylogenetic phylogenetic the in of Pachylepyrium of position couldanalyses, the when fall middle the of in added were Australia from specimens based mainly phylogeny on to the species (ReesEurasian et al. 2013). carbonicola be unambiguously determined 2015). et al. (Matheny into is subsumed genus Hygrometrici analyses that explicitly included the included the explicitly that analyses (and types Marasmius genus of the ) and sect. of sect.hence Marasmius Hygrometrici with clusters VOLUME 6 · NO. 2 Phylogenetic methods often allow, even in even often allow, methods Phylogenetic one formal than more trees, well-resolved A list of options should be 4. A list of options should options different discussed, final the for and arguments be tested, given. decision Weak support (or even absence of statistical statistical of even absence support (or Weak support) of proposed new genera indicates be cannot classifications alternative that rejected. is: advice so the ‘in dubiis And doubt refrain from in when abstine”, newproposing genera Example: warneri Lenzites Welti species, for this asa new it also name generic morphological discriminating does not have (2011) also & Hibbett did Justo characters. but not recognize either, genus it as a separate of concept their included it within The branching of the phylogenetic phylogenetic the of branching 3. The and sufficient should have trees support . statistical strong unsupported position within which is broader than that of Welti of Welti that than which is broader (2012). (ii) type the including species Only by (iii) of The position (i) The genus genus (i) The if the group in question is represented by is represented question group in if the Africa, species more tropical Asia, in many base should data the and Australia; and (c) include type of the included,being placement as the speciesdecides for a genus. to use which name Examples (iv) The type of Rubinoboletus CONCLUSIONS D (2015) Evolutionary relationships of polyphyly of the genus Psathyrella. Molecular Heimioporus and Boletellus (), with Phylogenetics and Evolution 46: 415–429. We encourage all mycologists undertaking an emphasis on Australian taxa including new Peintner U, Moser MM, Vilgalys R (2002) [“2001”] phylogenetic studies around the generic species and new combinations in Aureoboletus, Thaxterogaster is a taxonomic synonym of level to adhere to these guidelines, and Hemileccinum and . Australian Cortinarius: new combinations and new names. further invite editors and peer reviewers to Systematic Botany 28: 1–22. Mycotaxon 81: 177–184. bear them in mind when considering a work Hennig W (1950) Grundzüge einer Theorie der Petersen RH, Psurtseva N, Zmitrovich I, Chachuła MYCOLENS for publication. phylogenetischen Systematik. Berlin: Deutscher P, Arslanov S, Hughes KW (2015) Lignomyces, In recognizing that these problems Zentralverlag. a new genus of pleurotoid . are wide-ranging in mycology and may Hennig W (1965) Phylogenetic systematics. Annual Mycologia 107: 1045–1054. frustrate communication within the subject, Review of Entomology 10: 97–116. Redhead SA, Lutzoni F, Moncalv J-M, Vilgalys R we commend a cautious approach to Jenkinson TS, Perry BA, Schaefer RE, Desjardin DE (2002).Phylogeny of agarics: partial systematics introducing changes at generic rank. (2014) Cryptomarasmius gen. nov. established solutions for core omphalinoid genera in the Finally, we welcome and foresee critical in the Physalacriaceae to accommodate Agaricales (euagarics). Mycotaxon 83: 19–57. assessments of introductions of new taxa at members of Marasmius section Hygrometrici. Rees BJ, Midgley DJ, Marchant A, Perkins A, all taxonomical levels, especially families and Mycologia 106: 86–94. Orlovich DA (2013) Morphological and orders, in all groups of fungi. Justo A, Morgenstern I, Hallen-Adams HE, Hibbett molecular data for Australian Hebeloma species DS (2010) Convergent evolution of sequestrate do not support the generic status of Anamika. forms in Amanita under Mediterranean climate Mycologia 105: 1043–1058. REFERENCES conditions. Mycologia 102: 675–688. Sánchez-Garcia M, Matheny PB, Palfner G, Lodge Justo A, Hibbett DS (2011) Phylogenetic DJ (2014) Deconstructing the Baroni TJ, Kropp BR, Evenson VS, Wilhelm M classification ofTrametes (, (Agaricales) and introduction of the new genera (2014) Cercopemyces crocodilinus, a new genus ) based on a five-marker dataset. Albomagister, Corneriella, Pogonoloma and and species related to Ripartitella, is described Taxon 60: 1567–1583. Pseudotricholoma. Taxon 63: 993–1007. from North America. Mycologia 106: 785–796. Justo A (2014) The taxonomy of Turkey Tails Schoch CL, Seifert KA, Huhndorf S, Robert V, Bruns TD (2001) ITS reality. Inoculum 52 (6): 2–3. (Trametes) and related : One genus or Spouge JL, Levesque CA, Chen W, Fungal Buyck B, Hofstetter V, Eberhardt U, Verbeken A, too many? Mycophile 54 (4): 8–10. Barcoding Consortium (2012) Nuclear Kauff F (2008) Walking the thin line between Kirk PM (2015) Nomenclatural novelties. Index ribosomal internal transcribed spacer (ITS) Lactarius and Russula: the dilemma of Russula Fungorum 278. region as a universal DNA barcode marker for sect. Ochricompactae. Fungal Diversity 28: 15–40. Kluting KL, Baroni TJ, Bergemann SE (2014) Fungi. Proceedings of the National Academy of Co-David D, Langeveld D, Noordeloos ME (2009) Towards a stable classification of genera Sciences, USA 109: 6241–6246. Molecular phylogeny and spore evolution of within the Entolomataceae: a phylogenetic Tkalčec Z, Mešić A, Den C-Y, Pleše B, Ćetković H Entolomataceae. Persoonia 23: 147–176. re-evaluation of the Rhodocybe-Clitopilus clade. (2015) Hymenoporus paradoxus gen. et sp. nov., Dentinger BTM, Gaya E, O’Brien H, Suz LM, Mycologia 106: 1127–1142. a striking fungus of the family Omphalotaceae Lachlan R, Díaz-Valderrama JR, Koch RA, Kretzer A, Bruns TD (1997) Molecular revisitation (Agaricales, Basidiomycota) with tubular Aime MC (2015) Tales from the crypt: genome of the genus Gastrosuillus. Mycologia 89: hymenophore. Phytotaxa 204: 203–212. mining from fungarium specimens improves 586–589. Vellinga EC (2002) New combinations in resolution of the mushroom tree of life. Largent DL (1994) Entolomatoid fungi of the western Chlorophyllum. Mycotaxon 88: 415–417. Biological Journal of the Linnean Society: http:// United States and Alaska. Eureka, CA: Mad Vellinga EC, de Kok RPJ (2002) (1539) Proposal dx.doi.org/10.1111/bij.12553 River Press . to conserve the name Chlorophyllum Massee Eberhardt U, Verbeken A (2004) Sequestrate Lebel T, Tonkin JE (2007). Australasian species of against Endoptychum Czern. (). Lactarius species from tropical Africa: L. Macowanites are sequestrate species of Russula Taxon 51: 563–564. angiocarpus sp. nov. and L. dolichocaulis comb. (, Basidiomycota). Australian Vellinga EC, de Kok RPJ, Bruns TD (2003) nov. Mycological Research 108: 1042–1052. Systematic Botany 20: 355–381. Phylogeny and taxonomy of Macrolepiota Ge Z-W, Smith ME (2013) Phylogenetic analysis of Lodge DJ, Padamsee M, Matheny PB, Aime MC, (Agaricaceae). Mycologia 95: 442–456. rDNA sequences indicates that the sequestrate Cantrell SA et al. (2014) Molecular phylogeny, Vizzini A (2015) Nomenclatural novelties. Index Amogaster viridiglebus is derived from within morphology, pigment chemistry and ecology in Fungorum 244. the agaricoid genus Lepiota (Agaricaceae). (Agaricales). Fungal Diversity Welti S, Moreau OA, Favel A, Courtecuisse R, Haon Mycological Progress 12: 151–155. 64: 1–99. M, Navarro D, Taussac S, Lesage-Meessen L Gelardi M, Vizzini A, Ercole E, Horak E, Ming Z, Matheny PB, Moreau P-A, Vizzini A, Harrower (2012) Molecular phylogeny of Trametes and Li T-H (2015) Circumscription and taxonomic E, De Haan A, Contu M, Curti M (2015) related genera, and description of a new genus arrangement of Nigroboletus roseonigrescens gen. Crassisporium and Romagnesiella: two new Leiotrametes. Fungal Diversity 55: 47–64. et sp. nov., a new member of Boletaceae from genera of dark-spored Agaricales. Systematics Wu G, Feng B, Xu J, Zhu X-T, Li Y-C, Zeng N-K, tropical south-eastern China. PLOS One 10 (8): and Biodiversity 13: 28 –41. Hosen MI & Yang ZL (2014) Molecular e0134295. Nuhn ME, Binder M, Taylor AFS, Halling RE, phylogenetic analyses redefine seven major Greybeal A (1998) Is it better to add taxa or Hibbett DS (2013) Phylogenetic overview of clades and reveal 22 new generic clades in the characters to a difficult phylogenetic problem? the . Fungal Biology 117: 479–511. fungal family Boletaceae. Fungal Diversity 69: Systematic Biology 47: 9–17. Padamsee M, Matheny PB, Dentinger DTM, 93–115. Halling RE, Fechner N, Nuhn M, Osmundson McLaughlin DJ (2008). The mushroom T, Soytong K, Arora D, Binder M, Hibbett family Psathyrellaceae: evidence for large-scale

(68) IMA FUNGUS