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1 2 3 4 5 1 6 .. HIBBETT ,.BAUER ,.BINDER , A.. GIACHINI ,.HOSAKA ,A.JUSTO ,.LARSSON , 7 8 1,9 1 6 10 11 K.. LARSSON , J.D. LAWREY ,.MIETTINEN , .. NAGY , R.H. NILSSON ,M.WEISS , R.G. THORN

CONTENTS . ...... 396 G. ...... 397 I. Introduction ...... 373 H. ...... 399 A. Higher-Level Relationships ...... 374 I. ...... 400 . Taxonomic Characters and Ecological J. Jaapiales...... 402 Diversity...... 376 K. ...... 402 1. Septal Pore Ultrastructure ...... 376 L. ...... 403 2. Fruiting Bodies...... 380 M. ...... 405 3. Ecological Roles ...... 383 1. Atheliales and Lepidostromatales . . . . . 406 C. Fossils and Dating . . . . . 386 2. ...... 406 II. Phylogenetic Diversity ...... 387 3. ...... 407 A. ...... 387 4. ...... 409 B. ...... 389 III. Conclusions...... 411 C. ...... 390 References...... 412 D. ...... 391 1. ...... 391 2. ...... 392 3. ...... 393 I. Introduction 4. ...... 394 E. Trechisporales ...... 395 Agaricomycetes is a of that contains ca. 21,000 described , which is one-fifth of all known Fungi (Kirk 1Biology Department, Clark University, Worcester, MA, USA; e-mail: [email protected] et al. 2008). However, new taxa are continually 2Institut fu¨r Evolution und O¨ kologie, Universita¨tTu¨bingen, being described, and molecular ecologists rou- Auf der Morgenstelle 1, 72076 Tu¨bingen, Germany tinely detect DNA sequences of Agaricomycetes 3Centraalbureau voor Schimmelcultures, Uppsalalaan 8, that cannot be referred to known species, sug- Utrecht 3584 CT, Netherlands gesting that actual diversity of the group far 4Departamento de Microbiologia e Parasitologia, Centro de Ciencias Biologicas, Universidade Federal de Santa Catarina, exceeds the current catalog (Blackwell 2011; Caixa Postal 476, SC 88040 900 Florianopolis, Brazil Hibbett et al. 2011). Many members of Agari- 5Department of Botany, National Museum of Nature and comycetes produce conspicuous fruiting bodies Science, 4-1-1 Amakubo, Tsukuba, Ibaraki 305-0005, Japan that are popular subjects for artists and ama- 6 Department of Biological and Environmental Sciences, University teur naturalists (Petersen 2012). In addition, of Gothenburg, Box 100, 405 30 Gothenburg, Sweden most edible are Agaricomycetes, 7Natural History Museum, P.O. Box 1172 Blindern, 0318 Oslo, Norway including cultivated saprotrophs, such as Agar- 8Environmental Science and Policy Department, George icus bisporus (champignon), ostrea- Mason University, 4400 University Drive, Fairfax, VA, USA tus (oyster ), and edodes 9Botanical Museum, University of Helsinki, P.O. Box 7, 00014 (), and wild-collected ectomycorrhizal Helsinki, Finland (ECM) species, such as edulis (porcini), 10Fachbereich Biologie, Universita¨tTu¨bingen, Auf der Mor- genstelle 1, 72076 Tu¨bingen, Germany cibarius (), and Tricho- 11Department of Biology, University of Western Ontario, loma (matsutake). Psychoactive 1151 Richmond St. North, London, ON, Canada N6A 5B7 taxa, particularly species of , have

Systematics and Evolution, 2nd Edition The Mycota VII Part A D.J. McLaughlin and J.. Spatafora (Eds.) © Springer-Verlag Berlin Heidelberg 2014 374 D.S. Hibbett et al. been used both as recreational drugs and reli- parte (i.e., and - gious sacraments (Heim and Wasson 1958). ceae). Since 2007, three new orders of Agarico- Other members of Agaricomycetes are toxic, mycetes have been proposed: Amylocorticiales, with effects that range from gastrointestinal Jaapiales, and Lepidostromatales (Binder et al. distress, caused by diverse taxa such as Chlor- 2010; Hodkinson et al. 2013). This chapter pro- ophyllum molybdites, to life-threatening ama- vides a phylogenetic overview of Agaricomy- toxin poisoning, caused by phalloides, cetes, emphasizing recent molecular studies autumnalis, and others (Benjamin that address the diversity and phylogenetic 1995). The toxic compound phalloidin (from relationships of each (of course, A. phalloides) binds to actin, making it useful of Agaricomycetes classified as orders are sim- as a component of fluorescent stains for visua- ply mutually exclusive groups; they are not lizing the cytoskeleton. necessarily equivalent in age, number of spe- Agaricomycetes are not common as human cies, or phenotypic diversity). pathogens, although commune, which normally occurs as a wood-decay , is known to cause serious infections of lungs A. Higher-Level Relationships and other organs (Sigler et al. 1995). Several Agaricomycetes have been important as model All currently recognized orders of Agaricomy- systems in studies of fungal mating genetics cetes have been resolved as monophyletic in and development (S. commune, at least one analysis of rRNA , but sup- cinerea) (Ohm et al. 2010; Raper and Miles port for some groups has been weak or 1958; Stajich et al. 2010) and the absent, in part because of elevated rates of of wood decay ( chrysosporium, evolution in nuclear rRNA (nrRNA) genes in placenta, and others) (Martinez et al. certain Cantharellales and other lineages 2004, 2009). Finally, there is interest in uses of (Binder and Hibbett 2002; Binder et al. 2005; Agaricomycetes in industrial bioconversion Hibbett et al. 1997b; Moncalvo et al. 2006). processes and bioremediation (Ruiz-Duen˜as Genes encoding proteins, such as subunits 1 and Martı´nez 2009). and 2 of RNA polymerase II (rpb1, rpb2), Most of the taxa now classified in the Agar- mitochondrial ATPase subunit 6 (atp6), and icomycetes were included in a chapter on translation elongation factor 1-a (tef1), Homobasidiomycetes in the previous edition started to be used in fungal molecular system- of The Mycota (Hibbett and Thorn 2001). atics in the late twentieth century (Kretzer Eight informally named clades (e.g., euagarics and Bruns 1999; Liu et al. 1999; O’Donnell clade, russuloid clade) were proposed, based et al. 2001), and by 2006 a 6-, 200- almost entirely on analyses of ribosomal RNA species, -wide fungal phylogeny had (rRNA) gene sequences. A separate chapter been produced that included 37 species of treated (Wells and Ban- Agaricomycetes (James et al. 2006). The first doni 2001), which included jelly fungi and in-depth study of Agaricomycetes combining others with mostly septate basidia (Weiß et al. rRNA and protein-coding genes was that of 2004a). Today, Agaricomycetes is recognized as Matheny et al. (2007), who analyzed a 6.6 kb one of four major clades of , data set of rbp2, tef1, and nrRNA genes in 146 the others being the (see Ober- species (119 species of Agaricomycetes). This winkler 2014), (see Weiß et al. was the first analysis to provide strong sup- 2014), and (Fig. 14.1) port for the monophyly of Polyporales (which (Hibbett 2006; Padamsee et al. 2012). The 2007 had been weakly supported in rRNA ana- AFTOL classification of Fungi (Hibbett et al. lyses), and it suggested that the Sebacinales, 2007) included 17 orders of Agaricomycetes, Cantharellales, Auriculariales, and Phallomy- three of which contain species formerly classi- cetidae formed a paraphyletic assemblage, fied as Heterobasidiomycetes, namely Auricu- within which a clade containing the remain- lariales, Sebacinales, and Cantharellales pro ing Agaricomycetes is nested. Agaricomycetes 375

Agaricomycetidae AGARICOMYCOTINA AGARICOMYCETES

Phallomycetidae

DACRYMYCETES

TREMELLOMYCETES WALLEMIOMYCETES

Fig. 14.1 Higher-level phylogenetic relationships of Names in gray represent groups that have not been major groups of Agaricomycetes and other Basidiomy- included in phylogenomic analyses; placements of cota. The major topology (black lines and text) is based these taxa are based on studies combining rRNA on published Floudas et al. 2012; Padamsee et al. 2012) genes with 2–3 protein-coding genes (Hodkinson and unpublished (L. Nagy, D. Floudas, R. Riley, D. et al. 2013; Hosaka et al. 2006; Matheny et al. 2007) Hibbett et al., unpublished) phylogenomic analyses. 376 D.S. Hibbett et al.

Genome-based analyses are providing B. Taxonomic Characters and Ecological enhanced resolution and support for the Diversity higher-level relationships of Agaricomycetes, although so far only a few broad-scale phylo- Agaricomycete systematists have traditionally genomic studies of Agaricomycetes and other used morphological, biochemical, and ecologi- Fungi have been published (Hibbett et al. cal characters to formulate phylogenetic 2013). As of this writing, the most inclusive hypotheses and structure classifications, and a published analysis contains representatives of rich descriptive literature has evolved (Cle´men- 8 orders of Agaricomycetes (Floudas et al. c¸on 2004; Donk 1964;Ju¨lich 1981;Ku¨hner 1984; 2012), but over 70 Agaricomycete genomes Oberwinkler 1977; Petersen 1971a; Reijnders have been completed, many by the Joint and Stalpers 1992; Singer 1986). Nonmolecular Genome Institute of the US Department of characters that have been emphasized include Energy (Grigoriev et al. 2012). The phylogeny anatomical features (e.g., shapes and staining in Fig. 14.1 represents a consensus of pub- reactions of , basidia, and cystidia, lished (Binder et al. 2013; Floudas et al. 2012; hyphal systems of fruiting bodies, rhizomorph Padamsee et al. 2012) and unpublished (L. structures), macromorphology of fruiting bod- Nagy, D. Floudas, D. Hibbett, and R. Riley, ies (including developmental characters), pig- unpublished) phylogenomic analyses that col- ment chemistry, and cytological characters lectively draw on more than 40 whole-genome (e.g., nuclear behavior in basidiosporogenesis). sequences from 15 orders of Agaricomycetes, Cultural characters, wood-decay modes (white as well as representatives of Dacrymycetes, rot vs. brown rot), and asexual reproductive Tremellomycetes, Wallemiomycetes, and forms have also been used to address relation- other Fungi. Gomphales, Hysterangiales, Lepi- ships and provide tools for identification dostromatales, Phallales, Thelephorales, and (Nakasone 1990a; Redhead and Ginns 1985;Stal- Trechisporales have yet to be included in pers 1978). The previous version of this chapter phylogenomic analyses; placements of these (Hibbett and Thorn 2001) contained a review of groups in Fig. 14.1 are based on studies com- nonmolecular characters and ecological modes bining rRNA genes with protein-coding genes across major groups of Agaricomycetes, which is (Hodkinson et al. 2013; Hosaka et al. 2006; not repeated here. Part II of the present chapter Matheny et al. 2007). summarizes the major morphological and Phylogenomic analyses have confirmed ecological features within each order of Agarico- some aspects of the phylogeny of Agaricomy- mycetes, as informed by recent phylogenetic cetes that had been resolved in earlier studies of studies. The following sections discuss septal rRNA and protein-coding genes, such as the pore ultrastructure (Fig. 14.2), the evolution of monophyly of Agaricomycetidae (Agaricales, fruiting body forms (Figs. 14.3, 14.4, 14.5, 14.6, Boletales, Atheliales, Amylocorticiales, and 14.7, 14.8,and14.9), and the phylogenetic distri- Lepidostromatales) and its sister group rela- bution of major ecological modes (Table 14.1a, b) tionship to Russulales. Novel results from phy- across the major groups of Agaricomycetes. logenomics include the placements of Jaapiales, Corticiales, and Gloeophyllales. In previous 1. Septal Pore Ultrastructure analyses combining rRNA and protein-coding genes, was placed as the sister group to Septal pore ultrastructure provided clues to the Agaricomycetidae, and the higher-level posi- higher-level relationships of Agaricomycetes tion of Gloeophyllales was unresolved (Binder long before the advent of molecular characters. et al. 2010; Garcia-Sandoval et al. 2011). Recent The union of Dacrymycetes and Agaricomy- phylogenomic analyses indicate that Jaapiales, cetes is supported by their shared possession Gloeophyllales, and Corticiales form a strongly of dolipores that are surrounded at each side by supported clade, but its higher-level position is a more or less dome-shaped modified ER ambiguous. (endoplasmic reticulum) cisterna, the so-called Agaricomycetes 377

Fig. 14.2 Dolipores of Agaricomycetes in transverse (a–d, (b, c) sp. Note the imperforate SPCs in (b) f) or tangential sections (e) through the septal pore caps and the perforate SPCs in (c). (d, e) Schizophyllum com- (SPCs). Material was prepared by freeze substitution. mune. The regularly arranged perforations are especially ¼0.2 mmin(a)–(e), and 0.5 mmin(f). (a) Tulasnella sp. visible in (e). (f) sp. 378 D.S. Hibbett et al.

Fig. 14.3 Sebacinales (a, b), Auriculariales (c, d), and truncata.(e) tubaeformis.(f) cris- Cantharellales (e, f). (a) epigaea.(b) Cratero- tata. Photos by Michael Wood (a, e, f; http://www. colla cerasi.(c) gelatinosum.(d) mykoweb.com) and Jaroslav Maly (b–d) septal pore cap (SPC), also termed the par- this synopsis with new observations (by R. enthesome. In contrast, the SPCs of Tremello- Bauer) using serial sections. Accordingly, mycetes and Wallemiomycetes are composed of within the Dacrymycetes/Agaricomycetes saccules or fingerlike projections arising from union, four types of SPC are evident. the endoplasmic reticulum (SPCs are also occa- Imperforate to uni-perforate SPCs sionally absent in both groups) (Wells and Ban- (Fig. 14.2a). Probably depending on the sec- doni 2001; Padamsee et al. 2012). The tion, no or only one median perforation of ca. intracisternal space of the SPCs of Agaricomy- 100 nm appears in one section of the series. cetes is sandwiched by a fine electron-opaque This is typical for Dacrymycetes and layer so that the SPCs altogether appear nine- within Agaricomycetes for Sebacinales, Auri- lamellate in optimal sections (the double- culariales, Trechisporales, the sectioned cisternal membrane with three layers and Tulasnella clades within Cantharellales, each and the intracisternal lumen with three and the , , layers). Variation in the perforation of SPCs , Kneiffiella, and clades within Agaricomycetes was summarized by within Hymenochaetales [for the taxa and van Driel et al. (2009, and references therein). clades see Hibbett (2006) and van Driel et al. The following account completes and corrects (2009)]. Agaricomycetes 379

Fig. 14.4 Phallomycetidae, including Phallales (a), (c) coriaceum.(d) floccosus. Geastrales (b), Hysterangiales (c), and Gomphales (e) himantioides. Photos by Michael Wood (d, e). (a) ruber.(b) saccatum. (a–d; http://www.mykoweb.com) and Otto Miettinen (e)

Unstable perforate SPCs (Fig. 14.2b, c). In Regularly perforate SPCs with openings of all members of the Phallomycetidae studied, roughly 100 nm (Fig. 14.2d, e). Usually, the such as Aseroe sp., , Geastrum openings are more or less hexagonally spp., clavatus, impudicus, arranged. This type is realized in the Agarico- spp., and Sphaerobolus sp., there are mycetidae and related groups (i.e., Russulales, predominantly imperforate SPCs, but in each Corticiales, Gloeophyllales, Polyporales, and species, often in the same sections, there are Thelephorales), the and Peniophor- also dolipores with perforate SPCs. Usually, ella praetermissa clades within the Hymeno- the intracisternal layering of the imperforate chaetales, and the core cantharelloid group SPCs appears somehow incomplete. These within Cantharellales [for the taxa and clades observations suggest that the perforation for- see Hibbett (2006) and van Driel et al. (2009)]. mation in the Phallomycetidae begins later in Irregularly perforate SPCs with a few large comparison to that in the other accordant openings of several hundred nanometers groups. Accordingly, the SPCs in this group (Fig. 14.2f). This type characterizes the Cerato- may represent a unique type. basidiales clade within the Cantharellales. 380 D.S. Hibbett et al.

Fig. 14.5 Hymenochaetales (a–e) and Trechisporales sonii, with pustulate anamorphic regions. (g) (f–h). (a) sp. (b) Coltricia perennis;(c) hymenocystis.(h) Scytinopogon angulis- purpurea.(d) Rickenella fibula.(e) Xylo- porus. Photos by Otto Miettinen (a, b, e–g), Ellen Lars- don (¼Hyphodontia) crustosus.(f) Trechispora steven- son (c), Lasse Kosonen (d), and Nourou Yourou (h)

2. Fruiting Bodies ruber, Phallales) and veiled (e.g., lilacina, Agaricales), as well as rela- The morphological diversity of fruiting bodies tively simple corticioid forms (e.g., Trechispora in Agaricomycetes is unparalleled in any other stevensonii, Trechisporales). Agaricomycete clade of Fungi. Agaricomycete fruiting bodies fruiting bodies range over several orders of include complex, developmentally integrated magnitude in size, from tiny cyphelloid forms, such as stinkhorns (e.g., Clathrus forms, such as candidus or Agaricomycetes 381

Fig. 14.6 Polyporales. (a) sp. (b) ochroleuca. (c) femsjoeensis.(d) aurea. (e) sp. (f) sp. Photos by Otto Miettinen (a, b, d–f) and O. Manninen (c) the minute red algal parasite dil- (Agaricales) appearing seae (Agaricales), which are often less than and disappearing over a few hours, or peren- 1 mm in diameter (Binder et al. 2006; Boden- nial, with woody “conks” like fomentar- steiner et al. 2004), to giant , like ius (Polyporales) persisting for years. nobilissimus and ellip- The phylogenetic distribution of fruiting soideus (Hymenochaetales), which can be more body forms and hymenophore configurations than 1 m in diameter (Burdsall et al. 1996; Dai across orders of Agaricomycetes was reviewed and Cui 2011; Redberg et al. 2003). They may be by Hibbett (2007). All of the major morphotypes ephemeral, with deliquescent forms such as (e.g., pileate-stipitate, coralloid, polyporoid, 382 D.S. Hibbett et al.

Fig. 14.7 Gloeophyllales (a, b) and Thelephorales (c, d). (a) ponderosus.(b) fimbriata. (c) aurantiacum.(d) multiplex. Photos by Michael Wood (http://www.mykoweb.com)

Fig. 14.8 Russulales. (a) subcircellatus.(b) sulcata.(c) pyxidatus.(d, e). Macowa- nites vinaceodorus. Photos by Ellen Larsson (a–c) and M. Jeppson (d, e) corticioid, and gasteroid forms) have evolved and parsimony methods suggest that corticioid repeatedly. Several orders are composed exclu- forms may represent the plesiomorphic condi- sively (or almost exclusively) of resupinate tion in Agaricomycetes, with many indepen- forms (Amylocorticiales, Atheliales, Corticiales, dent origins of pileate-erect fruiting bodies, Jaapiales, Trechisporales, and Lepidostroma- as well as reversals to resupinate forms tales), and this is the only morphotype that is (Hibbett 2004; Hibbett and Binder 2002). known in all orders of Agaricomycetes (except These analyses should be interpreted with Geastrales, Hysterangiales, and Phallales). Ana- caution because they are based on limited lyses of character evolution using likelihood samples and simplistic models of Agaricomycetes 383

Fig. 14.9 Agaricomycetidae, including Agaricales (a, c), nus.(f) lacrymans. Photos by Guillermo Mun˜oz Amylocorticales (b), Atheliales (d), and Boletales (e, f). (a, c, e), Yu-Cheng Dai [b, reprinted from Binder et al. (a) .(b) bombycina.(c) Lyco- (2010), with permission of the Mycological Society of perdon perlatum.(d) salicum.(e) Boletus lupi- America], Paul Diederich (d), and Doris Haas (f)

3. Ecological Roles character evolution (Hibbett 2007). Nonethe- less, it is plausible that the ancestor of Agaricomycetes function as saprotrophs, the Agaricomycetes may have been a corti- pathogens, and mutualists. The group contains cioid fungus. Alternatively, the paraphyletic the major concentration of ECM taxa, as well as arrangement of Tremellomycetes and Dacry- white-rot and brown-rot decayers of massive mycetes within Agaricomycotina could indi- woody substrates. The distribution of cate that the ancestor was a . major ecological roles across the orders of 384 D.S. Hibbett et al.

Table 14.1a Nutritional modes in Agaricomycete orders, with examplar genera (Saprotrophsa)

White rot Brown rot Other/uncertainb Agaricales Pleurotus Coprinopsis Boletales Serpula Amylocorticiales Anomoporia Atheliales Athelopsis Lepidostromatales Polyporales Phanerochaete Postia Russulales Thelephorales Gloeophyllales Jaapiales Jaapia Corticiales Hymenochaetales Trichaptum Bridgeoporus? Trechisporales Porpomyces Hysterangiales Gomphales Kavinia Phallales Phallus Geastrales Sphaerobolus Auriculariales Sebacinales Cantharellales Botryobasidium aBold = genome sequence of at least one species completed; ? indicates uncertainty. Bacteriovores, algal and cyanobacterial parasites, and animal pathogens are not included. For references, see this text and Hibbett and Thorn (2001). Some endophyte observations by R. Gazis and R. Martin (unpublished) bThis broad category includes saprotrophs on soil, litter, dung, and keratinic substrates, as well as wood decayers of uncertain rot type or that do not correspond to classical white or brown rot definitions

Agaricomycetes is presented in Table 14.1a, b. talline cellulose occur in diverse lineages of Saprotrophic taxa occur in all orders of Agar- Agaricomycetes, and these may have retained icomycetes, and ECM taxa occur in at least 13 the plesiomorphic white-rot mode of saprotro- orders. As many as 37 independent ECM phy. Multiple origins of brown rot (in which lineages have been estimated to occur in Agar- lignin is not appreciably removed) in Polypor- icomycetes (Tedersoo et al. 2010; Rinaldi et al. ales, Boletales, and Gloeophyllales and the evo- 2008). It has been proposed that the ancestor of lution of the ECM condition in bicolor Agaricomycetes might have been ECM, based (Agaricales) seem to be associated with on the occurrence of ECM taxa in Sebacinales repeated losses of PODs and other PCW- (Weiß et al. 2004b), but molecular clock ana- degrading . Ongoing genomic compar- lyses (see below) suggest that the group is much isons are showing that some saprotrophic older than potential ECM hosts, including Pina- Agaricomycetes do not conform to the typical ceae (Floudas et al. 2012; Hibbett and Matheny models of either white rot or brown rot, includ- 2009). Phylogenomic analyses suggest that the ing wood decayers, such as Schizophyllum com- common ancestor of Agaricomycetes possessed mune and , and soil, litter, multiple ligninolytic II fungal peroxidases and dung fungi, such as bisporus and (PODs) and other plant cell wall (PCW)- Coprinopsis cinerea (Morin et al. 2012; Ohm decaying enzymes, implying that it was capable et al. 2010; Stajich et al. 2010). Similarly, of producing a white rot (in which both the increased sampling of ECM genomes is reveal- lignin and cellulose components of PCWs are ing considerable diversity in genes encoding degraded) (Floudas et al. 2012; Ruiz-Duenas PCW-degrading enzymes among indepen- et al. 2013). Wood decayers with multiple dently evolved symbiotic lineages (F. Martin PODs and enzymes capable of attacking crys- and colleagues, unpublished). Table 14.1b Nutritional modes in Agaricomycete orders, with examplar genera (Biotrophs)

Nematode- ECM Orchid Endophyte Mycoparasite Insect symbionts Lichenized Lichenicolous Bryophilous Plant pathogens trappers

Agaricales Laccaria Armillaria Pleurotus Boletales Amylocorticiales ? Atheliales Athelia Fibulorhizoctonia Atheliaarachnoidea Athelia Lepidostromatales Lepidostroma Phlebia Polyporales ? 385 Agaricomycetes Russulales Entomocorticium Thelephorales Gloeophyllales Jaapiales Corticiales Marchandiomphalina Hymenochaetales Coltricia Phellinus ? Rickenella Peniophorella Trechisporales Trechispora? “Trechispora ” Hysterangiales Hysterangium Gomphales Ramaria Phallales ? Geastrales Geastrum? Auriculariales Sebacinales Sebacina Cantharellales Cantharellus Tulasnella Sistotrema Burgella 386 D.S. Hibbett et al.

Environmental studies are expanding our C. Fossils and Molecular Clock Dating concepts of the ecological roles and diversity of Agaricomycetes (Hibbett et al. 2011). ECM and Molecular clock studies have yielded diverse soil communities have been studied intensively age estimates for Fungi, with the origin of the for many years (Horton and Bruns 2001; Peay Basidiomycota inferred to be anywhere from et al. 2008), but molecular environmental 450 million years ago (mya) to over 1 billion surveys are demonstrating the occurrence of years ago (Berbee and Taylor 2010; Blair 2009; diverse Agaricomycetes in other, often Douzery et al. 2004; Gueidan et al. 2011; Hedges surprising, habitats. For example, a small num- et al. 2004; Taylor and Berbee 2006). A genome- ber of freshwater, marine, and mangrove- based molecular clock analysis (Floudas et al. inhabiting Agaricomycetes are known from 2012) estimated the age of the Agaricomycetes cultures and fruiting bodies (Binder et al. at ca. 290 million years (with a 95 % highest 2006; Hibbett and Binder 2001; Frank et al. posterior density interval of 222–372 million 2010; Jones and Fell 2012; Yamaguchi et al. years). Other molecular clock studies using 2008), but recent studies using molecular rRNA genes, alone or in combination with approaches have detected Agaricomycetes in selected protein-coding genes, have focused marine planktonic communities (Gao et al. on groups within Agaricomycetes, such as Bole- 2010) and in corals, which seem to harbor spe- tales (Skrede et al. 2011; Wilson et al. 2012), cies of Agaricales, Auriculariales, Boletales, Agaricales (Matheny et al. 2009; Ryberg and Corticiales, Hymenochaetales, Polyporales, Matheny 2012), and brown-rot lineages and Russulales (Amend et al. 2012). The func- (Garcia-Sandoval et al. 2011). Taxon sampling tional biology of these marine taxa, known only in these analyses has been very divergent, and from DNA sequences, remains obscure. their results have often been inconsistent. For Numerous species of Agaricomycetes have example, an analysis focused on also been discovered as endophytes (Oses (Agaricales) (Matheny et al. 2009) suggested et al. 2008; Rungjindamai et al. 2008; Thomas that the group arose 143 (99–191) mya, while et al. 2008; Weiß et al. 2011). For example, another study that focused on Boletales but culture-based studies of the foliar and sapwood included diverse Agaricales (Skrede et al. endophytes of the rubber tree Hevea brasilien- 2011) suggested that the common ancestor of sis have detected many species of Polyporales Inocybaceae and existed ca. 45 (almost all white rot taxa), as well as Agaricales, (30–60) mya. Atheliales, Auriculariales, Cantharellales, New genome sequences are providing a Hymenochaetales, and Russulales (R. Gazis wealth of data for molecular clock analyses in and R. Martin, unpublished). Most of the sap- Agaricomycetes, but the paucity of reliably wood endophytes are closely related to known identified fossils continues to be a limiting fac- wood-decay species, suggesting that the endo- tor. Basidiomycetous hyphae with clamp con- phytes may exist as latent saprotrophs. Other nections are known from the Pennsylvanian ecological associations of Agaricomycetes that (ca. 330 mya) (Dennis 1970, 1976; Krings et al. have received significant attention recently 2011; see Taylor et al., Chap. 10, Vol. VII, Part include lichenized and lichenicolous forms B), but the earliest fossils that are clearly Agar- (DePriest et al. 2005; Diederich et al. 2011; icomycetes do not occur until the . Diederich and Lawrey 2007;Lawreyetal.2007) The oldest, Quatsinoporites cranhamii,isa and insect symbionts (Aanen et al. 2002; Mueller fragment of a poroid hymenophore from the et al. 2005;Nobreetal.2011; Slippers et al. 2003). lower Cretaceous (130–125 mya) that has sim- The latter group includes Fibulorhizoctonia ple (nonclamped) septate hyphae and hymenial (Atheliales), which produces sclerotia that elements that resemble setae, suggesting that it mimic the eggs of its symbionts may be a member of Hymenochaetales (Smith (Matsuura 2006;Matsuuraetal.2009). et al. 2004). Two gilled mushrooms that are Agaricomycetes 387 probably Agaricales are known from somewhat recognized species (Kirk et al. 2008), and is younger deposits, including represented on all continents. Effused, skinlike leggetti, from New Jersey (ca. 90– fruiting bodies characterize roughly half of the 94 mya) (Hibbett et al. 1995, 1997a), and species, e.g., in Sistotrema, Botryobasidium, Palaeoagaricites antiquus, from Burmese and Tulasnella, some of which are extremely amber (ca. 100 mya) (Poinar and Buckley delicate and inconspicuous. Stipitate-hydnoid 2007). Another fossil from , and stipitate-veined fruiting bodies occur in Palaeoclavaria burmites (Poinar and Brown the edible genera and Cantharellus, 2003), was originally interpreted as a clavarioid respectively, while coralloid fruiting structures member of the so-called (this are found in Clavulina and Multiclavula name refers to a polyphyletic taxon and is no (Fig. 14.3e, f). The hymenophore is mostly longer in use), but there are insufficient char- smooth but sometimes hydnoid or poroid, acters visible to determine its taxonomic place- while truly gilled structures are lacking. ment. Eocene fossils of Agaricomycetes include With the possible exception of the ectomycorrhizae associated with pine roots that Tulasnella (Rogers 1932), species in Cantharel- were interpreted as (Boletales) lales have a unique type of basidia called (LePage et al. 1997) and Appianoporites van- stichic, characterized by a longitudinal orienta- couverensis, a poroid fruiting body fragment tion of the spindle during meiosis, in contrast similar to . cranhamii (Smith et al. 2004). to the chiastic type with transversely oriented Dominican amber from the –Oligocene spindle present in all other Agaricomycetes. (ca. 15–30 mya) has yielded several well- While the presence of four-spored basidia con- preserved mushrooms that resemble extant stitutes an almost universal condition within Agaricales (Hibbett et al. 1997a, 2003; Poinar Agaricomycetes, it is not so in Cantharellales. and Singer 1990). Basidia with two sterigmata are found in, for Of the fossils listed previously, several have example, Clavulina and , and been repeatedly used as calibration points six or eight sterigmata predominate in Botryo- in molecular clock analyses, including Q. cran- and Sistotrema. Many species in hamii, A. leggetti,andtheputativesuilloid Cantharellus have predominantly five- (Floudas et al. 2012; Gueidan sterigmate basidia. The explanation for this et al. 2011;Skredeetal.2011). Additional fossils variation in sterigma number is not known, will surely be discovered, but it seems unlikely but a connection to the unique mode of meiosis that they will ever provide numerous rigorously is perhaps not unlikely. identified calibration points for the major clades Species in Cantharellales also show varia- of Agaricomycetes. Other sources of evidence tion in septal pore morphology. Botryobasi- that have the potential to address ages of diverse dium and Tulasnella are examples of genera lineages of Agaricomycetes include vicariant with imperforate parenthesomes, while species events and fossils of obligate symbionts, such in Cantharellus and Sistotrema and at least as ECM hosts (Hibbett 2001; Hibbett and some species in Ceratobasidium have perforate Matheny 2009; Matheny et al. 2009;Wilson parenthesomes (van Driel et al. 2009). et al. 2012) and the arthropods associated with Ecological diversity: most resupinate spe- taxa such as Termitomyces, ,and cies in Cantharellales seem to be saprotrophs. attine cultivars (Mikheyev et al. 2010;Nobre However, most species in Botryobasidium and et al. 2011;Slippersetal.2003). Tulasnella and the majority of species in Sisto- trema are capable of growing on common malt agar, a likely indication of the presence of cel- II. Phylogenetic Diversity lulolytic enzymes. On the other hand, no spe- cies in Cantharellales occur as primary A. Cantharellales decayers, and they do not develop an extensive within logs. Forthcoming genomes Overview: Cantharellales is a small order, com- of Botryobasidium botryosum, Sistotrema prising about 260 described and currently brinkmannii, Tulasnella calospora, and others 388 D.S. Hibbett et al. should provide clues to ecological capabilities (three genera) is characterized by a within the order. unique basidium morphology. The young basidium is Symbiotic relationships are widespread globose to club-shaped and develops four globose ste- rigma initials that at maturity become -shaped within the order and occur in all families before developing spores, which are forcibly dis- accepted here. ECM lineages include Canthar- charged. The unusual sterigmata have been interpreted ellus/Craterellus, Clavulina/Membranomyces, as monosporic epibasidia. All species can form second- Hydnum/Sistotrema sensu stricto, Ceratobasi- ary spores in the same way as described for Ceratoba- dium/Thanatephorus, and Tulasnella (Teder- sidiaceae. Hyphal septa have imperforate parenthesomes. Members of the form thin resu- soo et al. 2010 and references therein). pinate basidiomata or develop a loose mycelium within Another type of is present in the fruiting bodies of other resupinate fungi, apparently lichenized Multiclavula species that always without any interaction. Species seem to be saprotrophs grow associated with unicellular green algae or mutualists capable of forming orchid mycorrhizae (Lawrey et al. 2007). (Cruz et al. 2011; Preussing et al. 2010) or ectomycor- rhizae (Bidartondo et al. 2003). The nuclear ribosomal A parasitic lifestyle occurs in Ceratobasi- genes of large parts of the genus Tulasnella are inex- dium, where a common anamorph stage plicably deviant from those of other fungi and often known as is a widespread require tailored polymerase chain reaction primers for and troublesome crop pest, seemingly capable amplification (Taylor and McCormick 2008). of infecting a wide range of hosts (Mosquera- includes, as far as is known, a single genus, Botryobasidium. It is characterized by Espinosa et al. 2013; Parmeter 1970; Sneh et al. basidia that in most cases produce six or eight spores. 1996; Veldre et al. 2013). The anamorph genera A few species have spiny spores and four sterigmata. and Minimedusa, both related to Sisto- They were earlier referred to , but trema, are reported as parasites molecular data place all species examined firmly within (Diederich and Lawrey 2007). Botryobasidium (Binder et al. 2005). Secondary formation has not been observed in the family, and Systematics: the first comprehensive, septal pore parenthesomes are nonperforate. The basi- multiple-gene phylogeny of Cantharellales was diomata are very delicate, and hyphae are wide with a presented by Moncalvo et al. (2006). Veldre characteristic cruciate branching on subicular hyphae. et al. (2013) is the most recent phylogeny with Many species have an anamorph stage referred to the coverage of the whole order. These studies sup- form genus . They usually develop as separate, often brownish colonies sometimes port a division of Cantharellales into four integrated, however, with the teleomorph. Saprotrophy families, which may be defined by septal pore has been the assumed nutritional strategy, but a recent structure and secondary spore production. study detected orchid symbionts that, on the basis of DNA sequences, belong to Botryobasidium (Yukawa et al. 2009). There is no comprehensive molecular phy- Ceratobasidiaceae (eight genera) includes species with logeny for the family. thin, resupinate fruiting bodies developing on various (syn. , , kinds of fine woody debris and other plant remains, but Sistotremataceae; nine genera) is the largest family in also on living plants. Hyphae are broad and without Cantharellales in terms of the number of constituent clamps. With the exception of the type species of Cer- genera and the most diverse in terms of described atobasidium, all species studied so far have perforate species. As in many other cases, genera dominated by parenthesomes (van Driel et al. 2009; Weiß and Ober- corticioid species seem to represent the ancestral con- winkler 2001). Basidia are short, with 2–4 long sterig- dition, and lineages with erect fruiting bodies seem to mata. are capable of forming secondary have evolved from such species (Moncalvo et al. 2006). spores through the development of a functional ste- The corticioid species belong to Sistotrema and Mem- rigma from the primary spore. The formation of a branomyces. Sistotrema is a polyphyletic and ecologi- secondary spore has been interpreted as a second cally diverse genus. The type species forms a stipitate chance to send propagules into the air. This ability is fruiting body with a weakly hydnoid hymenophore and common also in Tulasnellaceae, Auriculariales, and is closely related to Hydnum. Other species related to Sebacinales but not known from other orders in Agar- the type have resupinate basidiomata with a poroid or icomycotina. A recent molecular study of Ceratobasi- hydnoid hymenophore. They all seem to form ectomy- diaceae suggests that only two or, perhaps, three genera corrhiza and share this strategy with the stipitate- should be recognized. Ceratobasidium is reduced to the hydnoid genus Hydnum (Nilsson et al. 2006). Membra- type species, and most other species are referred to nomyces largely shares the micromorphological char- Rhizoctonia (Oberwinkler et al. 2013a). acteristics, ECM habit, and phylogenetic placement Agaricomycetes 389 with the coralloid genus Clavulina. Cantharellus and to infundibuliform (Tremelloscypha) appear- Craterellus seem to make up a monophyletic group, ance. Sebacinalean basidiomes most often and together they form a third lineage within Hydna- have a gelatinous consistency. Known ana- ceae with ECM capacity. Fruiting bodies within this lineage are of the cantharelloid type, viz. more or less morphs in the Sebacinales comprise the species funnel-shaped and with a smooth, veined, or coarsely of Piriformospora, pycnidial conidiomata in semigilled hymenophore. Secondary spore production Craterocolla cerasi, and coremioid stages of is not known within this family, and the species exam- species of Efibulobasidium (Kirschner and ined have septa with perforate parenthesomes. Recent Oberwinkler 2009; Wells and Bandoni 2001). molecular studies in South America and Africa have unearthed a considerable number of new species and Sebacinales species have a worldwide distribu- lineages in the family (Buyck et al. 2013a, b; Henkel tion and are even known from Antarctica et al. 2011; Tibuhwa et al. 2012; Uehling et al. 2012a, b). (Newsham and Bridge 2010). A comprehensive However, a comprehensive phylogeny for the family is review of this group was recently published by still lacking. Several species in Hydnaceae (Cantharel- Oberwinkler et al. (2013b). lus cibarius, , and ) are highly prized as culinary mushrooms, Ecological diversity: over the past decade, yet attempts to keep these fungi in culture and to grow Sebacinales has received much attention mushrooms from those cultures have largely proved because of the exceptionally wide spectrum unsuccessful. As with Tulasnella, the nuclear ribosomal and the ubiquity of mutualistic associations genes of the genus Cantharellus are very deviant from with plant roots in which members of this those of other fungi (Moncalvo et al. 2006). group are involved. Sebacinalean mycobionts have been detected in ectomycorrhizae (Glen et al. 2002; Tedersoo and Smith 2013; Urban B. Sebacinales et al. 2003) and orchid mycorrhizae (Selosse et al. 2002; Warcup 1988), as well as in ericoid Overview: Sebacinales (Weiß et al. 2004b), one (Allen et al. 2003; Selosse et al. 2007), arbutoid of the basal clades in Agaricomycetes, presently (Hynson et al. 2013), and cavendishioid (Setaro includes 8 genera with ca. 30 described species et al. 2006) mycorrhizae, and even in junger- (Kirk et al. 2008). In contrast to these figures mannialean mycothalli (associations with liver- from the taxonomic literature, recent molecular worts) (Kottke et al. 2003). No other fungal phylogenetic studies have revealed a huge group is known to have a broader spectrum of amount of cryptic species in this group (e.g., mycorrhizal types. In addition, members of Riess et al. 2013; Selosse et al. 2007; Weiß et al. Sebacinales have recently been shown to occur 2011) and also suggest that generic concepts abundantly as endophytes in plant roots will have to be revised in the future to yield (Selosse et al. 2009; Weiß et al. 2011). A few monophyletic taxa. Morphological key features Sebacinales strains have been studied in vitro of Sebacinales include the ability of basidio- for their impact on plants in endophytic spores to form ballistoconidia (secondary associations. Most of these studies used the spores), dolipores with continuous parenthe- anamorphic strain Piriformospora indica and somes, longitudinally septate basidia, septa reported significant increases in growth and without clamp connections, and often thick- yield and improved resistance of the plant ened hyphal walls in substrate hyphae (Weiß hosts to abiotic and biotic stress (Qiang et al. et al. 2004b; Wells and Bandoni 2001; Wells and 2012), rendering the members of Sebacinales Oberwinkler 1982); unique apomorphies are promising bioagents for organic plant produc- not known for this group. There is a remarkable tion. The genome sequence of Piriformospora range of basidiome shapes in Sebacinales, from indica (Zuccaro et al. 2011) facilitates func- taxa that completely lack macroscopically visi- tional studies. Because of the richness of their ble basidiomes () to forms with cor- mutualistic associations with land plants, Seba- ticioid (Sebacina) (Fig. 14.3a), pustulate- cinales is a model group for studying the evolu- confluent (Efibulobasidium), cushion-shaped tion of plant–fungal interactions. (Craterocolla) (Fig. 14.3b), coralloid (Tremello- Though there is an increasing body of evi- dendron), or even stereoid (Tremellostereum) dence on the importance of Sebacinales as mutu- 390 D.S. Hibbett et al. alistic mycobionts of plant roots in terrestrial spectrum of basidiome shapes, including ecosystems, some species seem to have a sapro- effused (, ), effuso- trophic lifestyle (Craterocolla, Efibulobasidium). reflex (), odontoid (Stypella), hyd- Since P. indica and members of the morphospe- noid (Pseudohydnum)(Fig.14.3c), and infundi- cies Serendipita vermifera grow axenically in buliform (Tremiscus) basidiomes. Basidiomes of standard media, it can be assumed that many, some species even have a poroid or daedaleoid if not all, species of Sebacinales Group B (see habit (Elmerina [including Aporpium and Pro- below) have saprotrophic abilities. todaedalea], ) (Zhou and Dai Systematics: the monophyly of the Sebaci- 2013). All known species cause a white rot, nales has been demonstrated in molecular phy- some are regularly found on buried wood logenetic analyses (Weiß and Oberwinkler (Tremiscus helvelloides). Key characters of the 2001; Weiß et al. 2004b). All comprehensive Auriculariales include dolipores with continu- analyses of phylogenetic relationships within ous parenthesomes and the ability of basidios- Sebacinales have been based on nuclear- pores to form ballistoconidia (secondary encoded rRNA genes, including the internal basidiospores). With the exception of Hyaloria transcribed spacers (ITS) and partial large sub- pilacre, all species are ballistosporic. unit (nuc-lsu) regions. Most of the sequences Most of the known species in the Auricular- analyzed have come from environmental iales have longitudinally septate basidia, but sources; multilocus data derived from fruiting there are also species with transversely (Auri- bodies or cultures are needed to solidify the cularia) or obliquely septate (Patouillardina) systematics of Sebacinales. The available and even nonseptate (Oliveonia) or apically molecular phylogenetic analyses indicate that partially septate () basidia. Sebacinales is divided into two monophyletic In some genera (e.g., Myxarium, Protodontia, subgroups, informally known as Group A and Pseudohydnum, , Stypella, Tre- Group B. miscus), basidia have a plasma-devoid “stalk” (myxarioid, sphaeropedunculate basidia), Group A: Species forming macroscopically visible basi- which probably represents a taxonomically rel- diomes have only been reported from Group A.The evant character (Weiß and Oberwinkler 2001; types of interactions with plant roots are not uniformly Wells and Bandoni 2001). An explanation of distributed over Groups A and B; most of the reported this peculiar morphology was given by Bandoni taxa known to be involved in ectomycorrhizae belong to Group A, whereas sebacinalean mycobionts of ericoid (1984), who interpreted the basidial compart- mycorrhizae have only been reported from Group B. ments themselves as intrabasidial meiotic pro- Group B: The vast majority of Group B taxa are ducts (endospores) that in germination break known only from environmental sequences. S. vermi- the outer basidial wall and develop a single fera is the only known teleomorph in this group, yet it conidium, the “” in common ter- has been shown that this morphospecies is in fact a broad complex of cryptic species (Weiß et al. 2004b), all minology. In the myxarioid members of the of which may lack macroscopic basidiomes, that pro- Auriculariales the endospores do not fill the duce exceptionally long vermiform basidiospores and complete basidium but leave the characteristic are very poor in distinctive microscopic characters. It is stalk. possible that all teleomorphic species in Sebacinales Many investigated species show clamps Group B belong to this morphospecies and that the anamorphic genus Piriformospora, with two currently with characteristic retrorse projections (so- described species, evolved within this group from a S. called spurred clamps) (Bandoni and Wells vermifera-like ancestor that lost the ability to repro- 1992). Another characteristic microscopic fea- duce sexually. ture reported from many species of Auricular- iales is the ability of basidiospores to produce C. Auriculariales mostly crescent-shaped microconidia on short sterigmalike projections (Ingold 1982a, b). Overview: Auriculariales in its current concept Little is known about other anamorphs in includes ca. 30 genera with ca. 200 described Auriculariales. From recent reports about species (Kirk et al. 2008; Weiß et al. 2004a). It sporodochial, synnematous, bulbilliferous, and comprises wood-decaying fungi with a broad possibly also pycnidial examples in this group Agaricomycetes 391

(Kirschner 2010; Kirschner and Chen 2004; cola, Protomerulius, and possibly Tremello- Kirschner et al. 2010, 2012) we can extrapolate dendropsis; and a clade comprising that there might be a rich diversity of forms still Basidiodendron, Bourdotia, , and the to be detected. Their ecological function in cyphelloid bulbilliferous anamorph Ovipocu- nature is still unknown. lum (Weiß and Oberwinkler 2001; Zhou and Most species of Auriculariales are so-called Dai 2013). jelly fungi. As in Tremellomycetes, Dacrymy- cetes, and Sebacinales, the basidiomes of most members of Auriculariales have a gelatinous D. Phallomycetidae consistency and are able to experience drought conditions in a state of cryptobiosis, where the The group informally labeled the gomphoid- water content of the basidiomes is drastically phalloid clade (Hibbett and Thorn 2001) has reduced and the basidiomes revive and con- been classified as the subclass Phallomycetidae, tinue growing and sporulating when soaked with four orders: Geastrales, Phallales, Gom- again (Wells 1994). The Auricularia auricula- phales, and Hysterangiales (Hosaka et al. judae complex includes the broadly distributed 2006) (Fig. 14.4). wood-ear (mu-err) fungus, which occurs on dead wood and is one of the most important 1. Geastrales species of the world, particu- larly in Asia (Chang and Wasser 2012). The Overview: this group is represented by earth- group contains other edible mushrooms, for stars (Geastrum) (Fig. 14.4b), cannonball fungi example, Tremiscus helvelloides; however, (Sphaerobolus), and false truffles (, these are only sporadically collected in the Sclerogaster, and ). Taxa with nonse- field and not produced on an industrial scale. questrate fruit bodies possess an exoperidium Systematics: the monophyly of a core Aur- that opens in a stellate manner as it matures, iculariales (excluding and Exi- exposing the endoperidium with one (Geas- diopsis gloeophora) has been suggested by the trum) or multiple stomata () molecular phylogenetic analysis of Weiß and (Sunhede 1989). Most taxa, except Sclerogaster Oberwinkler (2001), who included the broadest and Sphaerobolus, have a brownish to blackish sampling of species of this group to date. Some , which becomes powdery at maturity. prior analyses using only rRNA genes focused Basidiospores of most taxa, including Scleroga- on Agaricomycetes (Binder et al. 2005; Hibbett ster, are globose with a warty to spiny ornamen- and Binder 2002) resolved the Auriculariales as tation. The fruiting body structure of a paraphyletic grade, but support for these Sphaerobolus is unique for Geastrales in having topologies has never been strong, and other a single peridiole instead of a powdery gleba. analyses show the group to be monophyletic. The mechanism of forcible ejection of peri- Multigene or phylogenomic analyses are still dioles was described in detail by Ingold (1972). lacking for this group. Given that many taxa Ecological diversity: the ecological charac- of Auriculariales have not yet been sequenced ters of this group have rarely been investigated. and the monophyly of the group is still tenta- Many species of the order grow on soil but tive, an infraordinal classification is not yet without obvious ECM plants nearby. In addi- available. Elements of a future classification tion, some species of Geastrum, Sclerogaster, may include a family, , com- and Sphaerobolus often fruit on rotten wood prising Auricularia, Eichleriella, Elmerina or wood chips (Hosaka and Castellano 2008), [including Aporpium and Protodaedalea], Exi- and Sphaerobolus fruits abundantly on artificial dia, Exidiopsis, and ; a clade com- media (Geml et al. 2005). Several species of prising Myxarium (including Hyaloria with Geastrum favor semiarid to arid environments, gasteroid sporulation) and the sporodochial for example, well-drained sandy soils of coasts anamorph Helicomyxa everhartioides; a clade and deserts (Kasuya et al. 2011). Such evidence including Heterochaetella, Protodontia picei- suggests that most, if not all, species in the 392 D.S. Hibbett et al. order are saprotrophic, as suggested by several Antarctica, should be separated into multiple authors (Kreisel 1969; Sunhede 1989). Most species. Therefore, a significantly higher trufflelike fungi are believed to form ectomy- number of species may be recognized in the corrhizae; Sclerogaster is an exception. One future. species of Geastrum, G. fimbriatum, has been described as forming ectomycorrhizae with Fagus (Agerer and Beenken 1998), but their 2. Phallales observation indicated the absence of a Hartig net. The ecological roles of Geastrales species Overview: this order is famous for its warrant further investigation. stinkhorns () and lattice stinkhorns Systematics: most taxa in Geastrales have (Clathraceae) (Fig. 14.4a), but recent molecular been treated in the order , along phylogenetic studies have shown that a number with () (Fischer 1900; of sequestrate taxa are also included. Most taxa Miller and Miller 1988; Zeller 1949), but early in the Phallales have fruiting bodies with a molecular studies (Hibbett et al. 1997a) demon- gelatinous layer and a gelatinous to mucilagi- strated that Geastrum and Lycoperdon, both of nous gleba. Fruiting bodies of epigeous stink- which possess a powdery gleba at maturity, are horns are often brightly colored (white and only distantly related. Kreisel (1969) first yellow to bright red) and composed of a pseu- segregated Geastrales from Lycoperdales but doparenchymatous receptacle with multiple did not provide a diagnosis. In addition, arms (Fig. 14.4a). The fruiting bodies of most Kreisel (1969) included only two genera, Geas- sequestrate taxa contain thick gelatinous layers, trum and Myriostoma, in the order. Hosaka and their gleba remains gelatinous to mucilagi- et al. (2006) formally described Geastrales nous. However, and Calvarula with a broader concept, including several pre- have a powdery gleba at maturity (Domı´nguez viously unrecognized taxa in the order. de Toledo and Castellano 1997). Spores of most Geastrales, which as a whole is moderately taxa are small, ellipsoid, smooth, and without supported as monophyletic, is divided into four ornamentation, but a few taxa, such as Kjeldse- families—Geastraceae, Sclerogastraceae, Sche- and Gastrosporium, have warty spore sur- nellaceae, and Sphaerobolaceae—that are all faces (Colgan et al. 1995; Domı´nguez de Toledo strongly supported as clades (Hosaka and and Castellano 1997). Castellano 2008; Hosaka et al. 2006). Geastra- Ecological diversity: most taxa are thought ceae, Sclerogastraceae, and Schenellaceae form to be saprotrophic due to their lignicolous a clade, with an ambiguous relationship among habit, but at least one species (Protubera canes- families (Hosaka and Castellano 2008). Within cens) has been reported to be ECM (Malajczuk Geastraceae, Myriostoma, the only taxon pos- 1988). This report, however, is suspect because sessing multiple stomata, represents the earliest P. canescens has recently been confirmed as an branch, suggesting that the evolutionary trend immature form of (May et al. 2010). is reduction from multiple stomata to a single It is likely that all members of the order are stoma. The early-diverging taxa within the saprotrophic, but further investigation is nec- order, Sphaerobolaceae and Schenellaceae, essary. form basidiospores in peridioles, and this may Phallales represents one of the prime exam- be the ancestral character state. ples of interactions of Fungi with arthropods A total of 7 genera and 64 species are cur- (Nouhra and Domı´nguez de Toledo 1994). The rently recorded in the order (Kirk et al. 2008), fruiting bodies of epigeous stinkhorns possess a but a number of undescribed species have been gleba that becomes slimy and malodorous at discovered for Geastrum and Sclerogaster maturity. The odor of a mature gleba attracts (Hosaka and Castellano 2008; Kasuya et al. a variety of mycophagous arthropods, espe- 2012). Furthermore, Kasuya et al. (2012) cially flies, that disperse the basidiospores demonstrated that Geastrum triplex, which (Tuno 1998). Unlike spores of many sequestrate was recorded from all continents except fungi, those of Phallales (including sequestrate Agaricomycetes 393 taxa) are rarely documented from mammal et al. 1999). They are characterized by a wide feces. It is possible that spore dispersal of Phal- range of fruiting body morphologies, from lales is entirely dependent on arthropods. stalked ramarioid/clavarioid (e.g., Ramaria, Systematics: Phallales was described by , and ) to club Fischer (1900) with two families, Phallaceae (), gilled (Gloeocantharel- and Clathraceae. A third family, Claustulaceae, lus), cantharelloid-gomphoid (Gomphus, was added to the order (Cunningham 1931; Phaeoclavulina, and Turbinellus) (Fig. 14.4d), Ju¨lich 1981; Zeller 1949), and this concept has tooth (), resupinate-odontoid (Kavi- been accepted for a long time. Miller and Miller nia) (Fig. 14.4e), all the way to sequestrate (1988) further expanded the ordinal concept by fungi (Gauteriaceae) (Giachini et al. 2010; Hos- including Protophallaceae in the order, but they aka et al. 2006; Humpert et al. 2001). also included (now in Hyster- Ecological diversity: members of Gom- angiales). Currently the order contains six phales show heterogeneity in their ecological families (Phallaceae, Clathraceae, Lysuraceae, characters. Most species in Beenakiaceae, Len- Protophallaceae, Claustulaceae, and - tariaceae, Kaviniaceae, , and ceae), and the monophyly of the order and Phaeoclavulina and some species of Ramaria each family is strongly supported by multigene (e.g., R. moelleriana, R. stricta, and R. circi- phylogenetic analyses (Hosaka et al. 2006). nans) grow and fruit on woody debris, a trait The basal grades of the order are composed that has led to their general categorization as of Protophallaceae, Claustulaceae, and Trap- saprotrophs. The other taxa of the order are peaceae, all of which exhibit an exclusively generally considered ECM, and while the nutri- sequestrate habit, indicating that stinkhornlike tional status of many species of Gomphales is fruit bodies are derived morphologies in Phal- still unknown, the formation of ectomycorrhi- lales (Hosaka et al. 2006). Among phylogenies zae by Turbinellus, Gomphus, and some of Agaricomycetes, Phallales represents the sole Ramaria species has been confirmed (Agerer example of an unambiguous transition from 1996a, b, c, d; Agerer and Iosifidou 2004; Agerer sequestrate to nonsequestrate forms. et al. 1998; Castellano 1988; Griffiths et al. 1991; A total of 29 genera and ca. 100 species are Masui 1926, 1927; Miller and Miller 1988; currently recorded in the order (Kirk et al. Nouhra et al. 2005; Rinaldi et al. 2008). 2008), but some genera, such as Protubera and Systematics: the and systematics Trappea, are polyphyletic, with species in both of the Gomphales has traditionally relied on Phallales and Hysterangiales (Hosaka et al. morphological characters now known to be 2006), and require further taxonomic revision. subject to parallel evolution and phenotypic Some new genera and species have been plasticity (Moncalvo et al. 2000). As a family, described recently (Cabral et al. 2012; Desjardin has traditionally been classified and Perry 2009). Because the center of diversity within the Aphyllophorales, along with dis- of this order probably lies in the tropics (Miller tantly related taxa such as Cantharellaceae, and Miller 1988) and many such areas have not , and (Donk been extensively investigated, the number of 1964). The phylogenetic relationships of mem- taxa in this group will be significantly higher bers of the order Gomphales, including its in the future. monophyly, have been estimated using mole- cular data (Giachini et al. 2010; Hosaka et al. 2006; Humpert et al. 2001), which revealed that 3. Gomphales gomphoid fungi are closely related to taxa in Geastrales, Hysterangiales, and Phallales Overview: the fungi in Gomphales (Ju¨lich 1981; in the subclass Phallomycetidae (Colgan et al. from the Greek pluglike) have long been recog- 1997; Giachini et al. 2010; Hibbett et al. 1997a; nized as a distinct, highly variable clade of Hosaka et al. 2006; Humpert et al. 2001; Agaricomycetes (Bruns et al. 1998; Hibbett Pine et al. 1999). Currently, the order encom- and Thorn 2001; Hosaka et al. 2006; Pine passes six well-supported families, namely 394 D.S. Hibbett et al.

Beenakiaceae (Beenakia, Kavinia and Ramari- exclusively sequestrate taxa with hypogeous cium), Clavariadelphaceae (), fruit bodies (e.g., Hysterangium, , Gautieriaceae (), Gomphaceae (Gloeo- ) (Fig. 14.4c), but taxa of epige- cantharellus, Gomphus, Phaeoclavulina, and ous habit (e.g., , Phallogaster) are also Turbinellus), Lentariaceae (Lentaria), and known, and they often expose a gleba at matu- Ramariaceae (Ramaria), distributed within 18 rity (Castellano and Beever 1994). Most taxa are genera and ca. 336 species (Kirk et al. 2008). characterized as having gelatinous to cartilagi- Despite their macromorphological varia- nous glebae of greenish to brownish tint, except tion, the members of the order share a number , which has a powdery gleba at of microscopic and macrochemical characters, maturity. including cyanophilic spore ornamentation, Ecological diversity: most taxa form ecto- chiastic basidia, similar hyphal construction, mycorrhizae with various host trees, including and positive hymenial reaction to ferric sulfate and in the Northern Hemi- (Donk 1961, 1964; Eriksson 1954; Humpert sphere and (mostly and et al. 2001; Petersen 1971b; Villegas et al. 1999, Leptospermum) and Nothofagaceae in the 2005). In the studies of Hosaka et al. (2006), Southern Hemisphere. In addition, some recent both Bayesian and parsimony analyses showed studies have extended the range of hosts to strong support for the monophyly of the Phal- Caesalpiniaceae, Phyllanthaceae, and Diptero- lomycetidae. Even though no definitive synapo- carpaceae (Castellano et al. 2000; Henkel et al. morphies have been identified for this 2012). Phallogastraceae is the only family in the gomphoid-phalloid clade, some potential syna- order with a saprotrophic habit. Some species pomorphic characters, including rhizomorph of Hysterangium form dense perennial hyphal morphology (presence of ampullate hyphae mats, which change the soil chemistry and and acanthohypha), pistillarin content, and microorganism biomass (Griffiths et al. 1994). structures of septal pore cap, have been pro- Fruiting bodies of Hysterangiaceae and Meso- posed (Agerer and Iosifidou 2004; Hibbett and phelliaceae are consumed by small mammals Thorn 2001). In addition, some members of the and marsupials, and they often make up a sig- gomphoid-phalloid clade, such as Gautieria, nificant portion of the animals’ diet (Claridge Hysterangium, Ramaria, and Geastrum, are 2002; Lehmkuhl et al. 2004). known to produce thick hyphal mats in soil Systematics: this order was proposed by (Agerer and Iosifidou 2004; Nouhra et al. Zeller (1939), whose treatment was followed 2005; Sunhede 1989). Although most of these by those of Locquin (1974) and Ju¨lich (1981). characters are not exclusive to the gomphoid- However, a Latin diagnosis was not provided phalloid fungi, the yellowish filled acanthocys- until Hosaka et al. (2006) formally described tidia and associated “exuded drops of pig- the order. The order is strongly supported as ments” have been reported only from the being monophyletic by multigene phylogenetic gomphoid-phalloid fungi [e.g., Geastrum, Gom- studies (Hosaka et al. 2006, 2008), but its rela- phus, Phallogaster, and Ramaria (Agerer and tionships to other orders in Phallomycetidae Iosifidou 2004)]. are not well supported. Hysterangiales is divided into four families, Hysterangiaceae, Mesophelliaceae, Gallacea- 4. Hysterangiales ceae, and Phallogastraceae, all of which are strongly supported as being monophyletic Overview: this group has long been considered (Hosaka et al. 2006, 2008). Hysterangiaceae a sequestrate (trufflelike) relative of stinkhorns and Mesophelliaceae form a clade, to which (Phallales). The ordinal status was accepted by Gallaceaceae is the sister family (Hosaka et al. some authors (Hosaka et al. 2006;Ju¨lich 1981; 2006). Phallogastraceae, the only saprotrophic Zeller 1939), but others have included the member of the order, represents the earliest group in the order Phallales (Kirk et al. 2008; branch, which is separated from the remaining Miller and Miller 1988). The order contains Hysterangiales (Hosaka et al. 2008), suggesting Agaricomycetes 395 that the ECM habit was gained only once within in the order form inconspicuous fruiting bodies the order. that rarely get collected and identified, the Fifteen genera and ca. 110 species are cur- known species number is likely a fraction of rently recorded in the order (Kirk et al. 2008), the true diversity. but a number of new species have recently been Fruiting body morphology ranges from discovered, mainly from the Southern Hemi- clavarioid (Scytinopogon), stipitate hydnoid sphere (Henkel et al. 2011; Hosaka et al. 2008). (Trechispora ), and resupinate poly- It has been demonstrated that many genera in poroid (Porpomyces, Trechispora) to corticioid this order are polyphyletic. For example, Hys- (Fig. 14.5f–h). Most species either have small terangium spp. are placed in both Hysterangia- spines (aculei) covering their hymenophore or ceae and Mesophelliaceae (Hosaka et al. 2008). are completely smooth. Fruiting-body-asso- Kirk et al. (2008) included Trappeaceae as the ciated rhizomorphs are common, and all spe- fourth family of the order, but because the cies have light-colored fruiting bodies that genus Trappea is also polyphyletic and the produce hyaline spores. Some dimitic species type species, Trappea darkeri, belongs to Phal- are found in Cristelloporia, , and lales (Hosaka et al. 2006), Trappeaceae should Trechispora, but most species are monomitic not be included in Hysterangiales. and bear clamps on all septa. Spore morphol- The biogeography of the order was exten- ogy is very variable, from very long and narrow sively studied by Hosaka et al. (2008), who spores of to tiny ellipsoid demonstrated that the ECM lineages (Hyster- spores of Porpomyces and spinose spores in angiaceae, Mesophelliaceae, and Gallaceaceae) most species of Trechispora. Conspicuous originated in the Southern Hemisphere (pre- subulate cystidia are found in Subulicystidium sumably east Gondwana), with a few range and . Calcium oxalate crystals are expansions to the Northern Hemisphere. common on subicular hyphae of Trechispora Although some area relationships can be and have been shown to be species-specific in explained by vicariance, many sister-group form (Larsson 1994). relationships, such as those of taxa from Aus- Ecological diversity: most species in the tralia and separated by short genus appear to be white-rot wood-inhabiting branches, can only be explained by long- (e.g., Sistotremastrum) or soil-inhabiting (e.g., distance dispersal, suggesting that trufflelike Porpomyces, Trechispora) saprotrophs. Trechi- fungi are capable of crossing ocean barriers. spora species are difficult to grow with standard culturing techniques for wood-decay fungi, whereas Sistotremastrum spp. pose no difficul- E. Trechisporales ties. Dunham et al. (2007) reported root- associated mycelial mats formed by Trechi- Overview: Trechisporales K. H. Larss. (2007) is spora, indicating a possible mycorrhizal associ- a relatively small order with ca. 100 species and ation, but further work is needed to confirm 8–13 genera. It was described only recently this inference. One species (misidentified as . (Hibbett et al. 2007), after DNA studies con- alnicola) has been reported as a grass parasite firmed it as a distinct clade (Binder et al. 2005; (Wilkinson 1987). Larsson 2004; Matheny et al. 2007). The major- Systematics: Larsson (2007b) divides the ity of species in the order belong to the genus order into two families: Ju¨lich Trechispora (including Cristelloporia, Scytino- 1982 (¼Subulicystidiaceae Ju¨lich 1982) with pogon), a highly diverse genus of mostly corti- Fibrodontia, , Porpomyces, Subulicysti- cioid fungi. The other genera contain only dium, Trechispora (including Cristelloporia, , with the exception of the Hydnodon), Tubulicium, and possibly Subuli- monotypic genus Porpomyces. A cium; and Sistotremastrum in its own, yet for- number of species in the order have an mally unnamed, family. Telleria et al. (2013) anamorphic stage: Aegerita tortuosa for produced a phylogeny of the order and Subulicystidium and Osteomorpha for confirmed that is also part Trechispora. Considering that almost all species of the Hydnodontaceae. Larsson et al. (2011) 396 D.S. Hibbett et al. and Birkebak et al. (2013) found that the cancer properties (Dai et al. 2010; Ju et al. 2010; clavarioid genus Scytinopogon was nested Wu et al. 2012). within Trechispora. Ecological diversity: the order exhibits a Only a few DNA-based species-level papers wide variety of different ecological strategies. on Trechisporales have been published: Albee- Most species of Hymenochaetales are white- Scott and Kropp (2010)onTrechispora, Telleria rot fungi. They are everywhere a major, and et al. (2012)onSistotremastrum, and Telleria often the dominant, part of the wood-rot com- et al. (2013)onBrevicellicium. Other genera munities (e.g., species of Hyphodontia, Phelli- that may belong to the order include the corti- nus, Trichaptum). Two polypore genera cioid Brevicellopsis, Dextrinocystis, and Dextri- (Coltricia and ) form ectomycorrhi- nodontia. A major open question in the zae (Tedersoo et al. 2007), and a number of systematics of the order is whether Trechispora species in other genera are parasites or patho- should be divided or kept together so that it gens of woody plants (e.g., many species of includes Cristelloporia, Echinotrema, Hydno- , Phellinus s.l., and ). Several don, and Scytinopogon, as most authors cur- species of Peniophorella have specialized rently do. organs for catching invertebrates, apparently an adaptation to a nitrogen-deprived environ- ment (Tzean and Liou 1993). A peculiar ecolog- F. Hymenochaetales ical group of mostly agarics are moss- associated. Whether the association is parasitic Overview: Hymenochaetales Oberw. 1977 is one or mutualistic is not clear (Larsson et al. 2006; of the larger orders of basidiomycetes, with Redhead 1981). over 900 species and ca. 75 currently recog- Systematics: Hymenochaetales as a clade is nized genera. The order is dominated by well supported, but its internal structure is wood-inhabiting polypores and bracket fungi largely unresolved. Larsson et al. (2006) (e.g., Phellinus, Trichaptum), as well as stereoid provided the only broad phylogenetic overview and corticioid fungi with a smooth or hydnoid of the order. They defined six clades using nuc- hymenophore (e.g., Hyphodontia, Resinicium). lsu rRNA sequences. Some of those clades were A few coralloid fungi (Alloclavaria, - not corroborated in other studies using nrDNA, chaete) and moss-associated agarics (e.g., Rick- and the branching order of the groups varies enella) are found in the order (Fig. 14.5a–e). from one analysis to another (Ghobad-Nejhad The largest known fruiting body belongs to and Dai 2010; Larsson 2007b; Miettinen and Phellinus ellipsoideus (Dai and Cui 2011). Larsson 2010). The genome of Fomitiporia Most Hymenochaetales species that have mediterranea has been published (Floudas been studied have dolipore septa with continu- et al. 2012), and two additional genomes (Rick- ous (imperforate) parenthesomes, in contrast enella mellea, ) were to most other polypores, agarics, and corticioid produced in the US Department of Energy fungi (van Driel et al. 2009). Peniophorella prae- Joint Genome Institute in 2013. termissa, a corticioid fungus, is the only species in the order reported with perforate parenthe- somes, but much of the diversity in the order Hymenochaetaceae: this family contains 60 % of remains unstudied in this respect. described Hymenochaetales species, mostly poly- pores, a number of stereoid fungi, and a few hydnoid Economically important pathogens of trees fungi. All species in this family are characterized by include in rubber and brown pigments that turn black in KOH (xantho- other tropical tree plantations (Farid et al. chroic reaction). Hyphae are simple-septate and par- 2009) and on temper- enthesomes imperforate in all species studied so far. ate conifers (Lim et al. 2005). The fruiting Many species have characteristic brown cystidia, setae, in the (which explains the name of bodies of Inonotus sanghuan, many Phellinus the type genus, Hymenochaete). Due to morphologi- spp., and cankers of I. obliquus are used in cal similarities, this order has long been recognized herbal medicine and are reported to have anti- in the literature in a way that corresponds to the Agaricomycetes 397 current concept: Phellinus s.l. (dimitic polypores), Oxyporus clade: this small clade contains the Inonotus s.l. (monomitic polypores), Hymenochaete polypore genus Oxyporus and poroid-hydnoid Botryo- s.l. (corticioid and hydnoid), and (hyd- dontia (Sell et al. 2013). The extent of the genus noid). All species are wood inhabiting, and many Oxyporus and delimitation against Rigidoporus is species are parasites of living trees and bushes. A unclear. strange exception is parasitica, which Rickenella clade: this is the most diverse in reportedly grows on living leaves (Wagner and terms of morphology and ecological strategies of all Ryvarden 2002). Whether the ECM Coltricia and Col- the Hymenochaetales clades and includes moss- triciella, traditionally assigned to Hymenochaetaceae, associated agarics (e.g., Rickenella), stereoid (Cotyli- belong here is unclear (see below). Recent regional dia), clavarioid (Alloclavaria), poroid (), and morphology-based treatments of Hymenochaetaceae many wood-rotting corticioid (Peniophorella, Resini- include Nun˜ez and Ryvarden (2000) for East Asia, cium) species. Larsson (2007b) and Miettinen and Ryvarden (2004) for the Neotropics, and Dai (2010) Larsson (2010) found this clade to be paraphyletic, for China. The basis for DNA-based assessments of and clearly multigene data sets are needed to resolve generic concepts in the group comes from studies of the structure in this part of the fungal tree. It may be Wagner and Fischer (2001, 2002a, b). Later family- basal to the order, but again the current DNA data do level studies have been few and based on one or two not permit strong statements in this regard. A hand- ribosomal genes only (Jeong et al. 2005; Larsson et al. ful of studies have been conducted at the genus level 2006; Parmasto et al. 2013). Many of the newly in this group: Moncalvo et al. (2002) and Redhead defined genera have received attention in more et al. (2002) dealt with agarics, Dentinger and focused phylogeny papers: Inonotus (Tian et al. McLaughlin (2006) with Alloclavaria, Sjo¨kvist et al. 2013), Phellinus s.s. (Cui and Decock 2012; Decock (2012) with and Muscinupta, Larsson et al. 2006; Fischer and Binder 2004; Tomsˇovsky´ et al. (2007b) with Peniophorella and other s. 2010b), (Zhou and Qin 2013), l., Miettinen and Larsson (2010) with Sidera, and (Keller and Hohn 1997), (Brazee and Nakasone (2007, 2012) with Resinicium and Tsugacor- Lindner 2013; Tomsˇovsky´ et al. 2009), and Phyllo- ticium. Hallenberg et al. (2007) studied the Penio- poria (Valenzuela et al. 2010; Wagner and Ryvarden phorella praetermissa species complex. 2002; Zhou and Dai 2012). The genus Fomitiporia in Lineages of uncertain position: Coltricia and particular has been the subject of many studies Coltriciella are the only ECM genera in the order. (Amalfi and Decock 2013; Amalfi et al. 2010, 2012; Larsson et al. (2006) included them in the same Decock et al. 2005, 2007; Fischer 2002; Fischer et al. clade (named Coltricia clade) with parts of Hypho- 2005; Vlasa´k and Kout 2010). The corticioid genus dontia (the here). The nrDNA of Col- Hymenochaete has turned out to be polyphyletic, and tricia is highly divergent from that of other part of the species belongs to a new genus, Pseudo- Hymenochaetales and occupies a long branch in chaete (He and Dai 2012; He and Li 2013; Parmasto nrDNA-based analysis, jumping around in phyloge- et al. 2013; Wagner and Fischer 2002a). The genus nies. Considering the long branch and divergent ecol- now also includes poroid spp. and hyd- ogy and morphology, the position of Coltricia and noid spp. The position of the Hymeno- Coltriciella within the order is in need of further chaete-like coralloid genus Clavariachaete has not study. Tedersoo et al. (2007) provide the only DNA- been studied, but morphologically it is a typical based study of the group. The wood-rotting polypore member of the Hymenochaetaceae (Parmasto 2010). and corticioid genera Trichaptum, , Schizoporaceae: this species-rich clade contains the Cyanotrama, , and Poriodontia belong in bulk of corticioid Hyphodontia, now classified in the the vicinity of Hymenochaetaceae, but not within it. genus , one of the largest genera of wood-rotting These genera seem to be somewhat closely related; fungi (Hjortstam and Ryvarden 2007, 2009). Most poroid they do not seem to form a monophyletic group Hyphodontia or Schizopora belong here, and micromor- (Binder et al. 2005; Ghobad-Nejhad and Dai 2010; phologically the clade is relatively homogeneous. Larsson Larsson et al. 2006; Miettinen and Larsson 2010). et al. (2006) included the genus Coltricia here and The polypore genus Bridgeoporus is a segregate of called it the Coltricia clade. Here we consider the position Oxyporus and has been shown to belong to Hymeno- of Coltricia unresolved. No phylogenetic overview of chaetales (Redberg et al. 2003). It does not seem to be Hyphodontia s.l. or this clade exists. Paulus et al. (2000) very closely related to Oxyporus, and its position in studied species phylogeny in Schizopora. the order is open. Tubulicrinacae: initially called the Hyphodontia clade, Larsson (2007b) used the family name Tubulicrinaceae for this clade. It contains inconspicuous G. Polyporales wood-rotting corticioid fungi with a variable micromor- phology. The largest genus is Tubulicrinis. Overview: Polyporales (¨umann 1926) Kneiffiella clade: Kneiffiella is another segregate genus of Hyphodontia sensu lato with many hydnoid includes approximately 1,800 described species species. Most species in the clade have characteristic (Kirk et al. 2008), making it one of the larger tubular tramal cystidia. orders of Agaricomycetes. However, new spe- 398 D.S. Hibbett et al. cies are continually being described, even in are the model systems for white-rot and brown- relatively well-studied areas such as western rot biochemistry (respectively), are both in the Europe (Bernicchia et al. 2010; Spirin et al. Polyporales (Martinez et al. 2004, 2009). No 2012; Vampola and Vlasak 2012), and the num- mycorrhizal taxa are known in the order. ber of species known only from environmental Many members of Polyporales are commonly sampling and studies of endophytic commu- isolated as part of the endophytic communities nities has increased dramatically in recent in woody tissues and roots, and though several years (Fro¨hlich-Nowoisky et al. 2009, 2012; ecological roles have been proposed for these Hallenberg et al. 2008). Polyporales contains fungi, from latent saprotrophs to protective conspicuous bracket fungi, including perennial agents, their true function remains largely “conks” (e.g., applanatum, Fomes unknown (Porras-Alfaro et al. 2011). fomentarius), as well as more cryptic effused Systematics: approximately 150 genera and (resupinate) forms, which often fruit on the 40 legitimate family names are available for use undersides of logs (Fig. 14.6b–d). Other species in Polyporales (Larsson 2007b; Ryvarden 1991), have pileate-stipitate fruiting bodies or multi- but there is no broadly accepted consensus ple flabelliform lobes (e.g., , Hydnopo- infraordinal classification. Recent monographs lyporus) (Fig. 14.6a, e, f). The hymenophore is on Polyporales include those of Nun˜ez and frequently poroid (e.g., Polyporus) but can also Ryvarden (2000) on East Asian polypores, be hydnoid (), lamellate (Tra- Ryvarden (2004) on neotropical polypores metes [Lenzites] betulina), merulioid (Phlebia), (Ganodermataceae), Niemela¨ (2005)and or smooth (Phanerochaete). No gasteroid taxa Bernicchia (2005)onEuropeanpolypores,and are known, but tigrinus has a naturally Bernicchia et al. (2010) on European corticioid occurring form in addition to the fungi. typical agaricoid form (Hibbett et al. 1994). A The monophyly of Polyporales was not well few species produce underground sclerotia supported in analyses of rRNA gene sequences (e.g., , Polyporus, Wolfiporia). The (Binder et al. 2005; Larsson 2007b). However, order has varied hyphal anatomy, including analyses adding single-copy protein-coding monomitic forms (with only generative hyphae, genes (Garcia-Sandoval et al. 2011; Justo and e.g., ), as well as dimitic and trimitic Hibbett 2011; Matheny et al. 2007; Miettinen forms (with thick-walled skeletal or binding et al. 2012; Sjo¨kvist et al. 2012) and genome- hyphae) (Gilbertson and Ryvarden 1986). No based phylogenetic analyses (Binder et al. 2013; morphological synapomorphy characterizes Floudas et al. 2012) have strongly supported the the Polyporales, and the most common mor- monophyly of the order. The sister group of phological types described previously also Polyporales is not known with confidence; in occur in other orders of Agaricomycetes. the studies just mentioned, Corticiales, Gloeo- Ecological diversity: along with phyllales, Russulales, and Thelephorales usu- members of Hymenochaetales and Russulales, ally seem to be closely related to Polyporales, members of this order dominate wood-decay but relationships between these orders remain communities in terrestrial ecosystems. A few in need of further study. species act as plant pathogens, causing timber The studies of Binder et al. (2005, 2013) damage (e.g., species of Ganoderma, Fomitop- provide broad overviews of the major lineages sis, and Wolfiporia), and others are major decay of Polyporales based on taxon-rich sampling of agents of structural timber (e.g., ). rRNA genes in combination with rpb1, rpb2, Wood decayers in Polyporales can be divided and tef1 sequences, as well as gene-dense phy- into two major groups: white-rot species, which logenomic analyses. Four major groups have are able to decay both lignin and cellulosic been informally labeled as the Antrodia, core compounds, and brown-rot species, which polyporoid, phlebioid, and “residual” clades, remove cellulose and hemicellulose without sig- the latter being more a mixed bag of taxa that nificant lignin degradation (Worrall et al. did not fit in the other clades. Larsson (2007b) 1997). P. chrysosporium and P. placenta, which and Miettinen et al. (2012) sought to apply Agaricomycetes 399 existing family names to parts of the clades taxa (Lentinus) are also nested in the clade. This is recognized in phylogenetic analyses. the best sampled lineage of Polyporales, both in terms of taxa and genes, and the only one with a well- supported internal structure. Three major lineages Residual polyporoid clade: the monophyly of this clade were recognized by Justo and Hibbett (2011), termed remains uncertain. All taxa in this group produce a the , trametoid, and Polyporus clades. white rot, which is probably plesiomorphic for the Representative genera with recent phylogenetic studies Polyporales (Floudas et al. 2012), but morphologically include Lentinus (Grand et al. 2010), they are very diverse and include poroid (Rigidoporus), (Li and Cui 2013), (Cui et al. 2011), agaricoid (Panus), corticioid (Hyphoderma), Perenniporia s. lato (Decock and Ryvarden 2003; resupinate-hydnoid (Steccherinum), and stipitate- Robledo et al. 2009; Zhao et al. 2013), Polyporus s. lato steroid forms (Podoscypha). Representative taxa that (Kru¨ger 2008, 2010; Kru¨ger and Gargas 2004; Sotome have been the subject of recent phylogenetic studies et al. 2008, 2013), and (Justo and Hibbett include (Miettinen et al. 2012), Cerrena 2011; Tomsˇovsky´ 2008; Welti et al. 2012). (Lee and Lim 2009), Hyphoderma (Larsson 2007a), Lineages of uncertain position: three relatively (Telleria et al. 2010), Pseudolagarobasi- small lineages of white-rot polypores seem to be closely dium (Hallenberg et al. 2008), Podoscypha (Sjo¨kvist related to the Antrodia or core polyporoid clades, but et al. 2012), and Steccherinum (Miettinen et al. 2012). they apparently do not belong to either group (Binder The group exemplifies the numerous transitions in et al. 2013; Miettinen and Rajchenberg 2011). These hymenophore types and microscopic characters (e.g., lineages include the genus , the cystidia and hyphal types) that have occurred repeat- clade (, , Tyromyces), and the edly during the evolution of the Polyporales (Miettinen / clade (Cinereomyces, Gelato- et al. 2012). poria, , Sebipora). Resolving the position of these Phlebioid clade: largely dominated by corticioid lineages and the phylogenetic structure of the residual forms, much of the taxonomy of this diverse group polyporoid clade are two major issues for the higher- revolves around two large, highly polyphyletic genera, level taxonomy of Polyporales. Improving the internal Phlebia and Phanerochaete, and their limits and rela- resolution in the phlebioid, antrodia, and core polypor- tions with respect to several smaller genera. A number oid clades is necessary to move forward in the family- of polypore genera are also found in the clade (e.g., level and generic taxonomy of these groups. , Ceriporia and ). Taxa that have been the subject of phylogenetic studies include Ceri- poria (Jia et al. 2013), (Tomsˇovsky´ et al. 2010a), Phanerochaete (De Koker et al. 2003; Greslebin H. Thelephorales 2004; Wu et al. 2010), and (Tomsˇovsky´ 2008). , a close relative of Ceriporia,is Overview: Thelephorales is a strongly sup- often considered a brown-rot fungus (Gilbertson and ported clade that currently includes ca. 18 Ryvarden 1986) and would be the only brown rotter in genera and 269 described species (Kirk et al. this lineage of white-rot taxa (Lindner and Banik 2008). 2008). The group is morphologically diverse Antrodia clade: this lineage includes exclusively species that produce a brown-rot type of decay. The and contains corticioid (Tomentella), canthar- majority of all known brown-rot fungi belong to this elloid (Polyozellus multiplex), clavarioid (The- clade. Pileate and resupinate polypores are predomi- lephora), and pileate forms (Hydnellum) nant, along with a few corticioid taxa (e.g., , (Fig. 14.7c, d). Hymenophores of pileate taxa possibly ). Several genera have received may be poroid (), toothed (Hydnel- attention in phylogenetic studies, including Antrodia sensu lato (Bernicchia et al. 2010; Rajchenberg et al. lum, ), smooth to wrinkled or tuber- 2011; Spirin et al. 2013; Yu et al. 2010), culate (Thelephora), or lamellate (Lenzitopsis). (Lindner et al. 2011), (Kim et al. 2007), It was once suggested that the pileate-stipitate (Lindner and Banik 2008), Postia sensu Horakia (¼) was related to lato (Pildain and Rajchenberg 2013), and Sparassis based on spore morphology (Dai et al. 2006; Wang et al. 2004). A general overview of the clade is given by Ortiz-Santana et al. (2013), who (Oberwinkler 1975), but molecular data place showed that generic delimitation remains highly prob- it in Agaricales (Matheny et al. 2006), as had lematic, with most of the traditionally recognized been suggested by Singer (1986). Basidiospores genera being poly- or paraphyletic. are mostly dark, ornamented, and with a Core polyporoid clade: this group roughly corre- distinctive angular outline but may also be sub- sponds to the families Polyporaceae and Ganoderma- taceae in the sense of Ryvarden (1991) and includes globose and spinose ( and ). mostly polypores with a trimitic hyphal system. Thelephoric acid (a terphenyl quinone, simi- Some corticioid (, ) and agaricoid lar to atrotomentin) is found in Bankera, 400 D.S. Hibbett et al.

Boletopsis, Hydnellum, Phellodon, Polyozellus, netic studies and included previously in the , Sarcodon, and Thelephora. Vuilleminiales (Boidin et al. 1998), the Dendro- This compound also occurs in Boletales and clade (Binder and Hibbett 2002), and other orders but nonetheless seems to be a the corticioid clade (Binder et al. 2005; Larsson distinguishing feature of the group (Bresinsky et al. 2004). It has been represented by a single and Rennschmid 1971). family, Corticiaeae Herter, made up of ca. 29 Ecological diversity: most members of The- genera and 136 species (Kirk et al. 2008), but lephorales are ECM and are often dominant the molecular phylogenetic study of Ghobad- components of mycorrhizal communities Nejhad et al. (2010) recognized three families (Bruns et al. 1998; Tedersoo et al. 2010). How- and several new genera (discussed subse- ever, Lenzitopsis produces fruiting bodies on quently). The order is made up of mostly resu- wood of junipers and is reported to produce a pinate species that produce smooth white rot (Zhou and Ko˜ljalg 2013). hymenophores, a monomitic hyphal system is also reported to grow on wood of living trees with or without clamps, and smooth basidios- (U. Ko˜ljalg, unpublished) and is presumably pores, often with pink walls. Many species pro- nonmycorrhizal. Amaurodon has been placed duce pink or red basidiomata with a as the sister group to the remaining Thelephor- catahymenium in which young basidia do not ales, but Lenzitopsis is nested within the group, form a palisade but are formed deep within a closely related to Tomentellopsis, suggesting layer of hyphidia and then elongate to reach the that there have been multiple transitions hymenial surface, and some have dendrohyphi- between nutritional modes (Larsson 2007b; dia; however, there is no morphological synap- Zhou and Ko˜ljalg 2013). omorphy that characterizes the entire order. Systematics: the current classification of Some species are known only from asexual Thelephorales (Kirk et al. 2008) includes two stages. One genus of these is the anamorph- families, Thelephoraceae and . typified Marchandiomyces, which was com- Donk (1964, p. 247) thought that the similarity monly included as a core genus in recent of the Bankeraceae to certain Thelephoraceae molecular studies. The genus was originally was “an example of extreme convergence,” but established for asexual lichen parasites (Dieder- other authors suggested that the two families ich 1990; Etayo and Diederich 1996), but sexual were closely related (Ju¨lich 1981; Stalpers 1993), forms are also now known for the group. These and this has been repeatedly supported by include Marchandiobasidium aurantiacum as molecular data (Binder et al. 2005; Bruns et al. the teleomorph of Marchandiomyces aurantia- 1998; Larsson 2004, 2007b; Zhou and Ko˜ljalg cus (Diederich et al. 2003) and Marchandiopsis 2013). The analysis of Zhou and Ko˜ljalg (2013) quercina, a species previously assigned to Lae- resolved two nonsister clades corresponding ticorticium or that was found to be to Bankeraceae (one including Bankera and nested among asexual Marchandiomyces spe- Phellodon and another containing Hydnellum, cies by Ghobad-Nejhad et al. (2010). The clade Sarcodon, and Boletopsis) and a paraphyletic containing Marchandiomyces also includes assemblage of taxa corresponding to Thele- described plant pathogens in the teleomorph- phoraceae. However, internal support for typified genera and , many deep nodes was weak. An in-depth multi- indicating that sexual–asexual relationships gene phylogenetic analysis is needed to assess among these species will require more study. the classification of the order. The type species of the order, Corticium roseum, may form a bulbil-like anamorph known as Hyphelia rosea; these bulbils are sim- I. Corticiales ilar to those of Laetisaria and Marchandio- myces (Eriksson and Ryvarden 1976). Overview: Corticiales K.H. Larsson is a small Ecological diversity: fungi in Corticiales order established to accommodate basidiomy- exhibit a remarkable range of ecologies, cetes recognized in recent molecular phyloge- including saprotrophs, plant pathogens, lichen Agaricomycetes 401 pathogens, and lichenized species (Lawrey et al. 2002; Larsson 2007b; Larsson et al. 2004; 2008). The basal position of saprotrophic spe- Lawrey et al. 2008). The sister group of cies in Vuilleminiaceae and Punctulariaceae Corticiales seems, on the basis of many of would indicate that this is the ancestral condi- these studies, to be the order Gloeophyllales. tion for the order, but the most derived clade, One species of Corticiales, Punctularia , contains a complex mixture of strigoso-zonata, has been subject to whole- ecological forms, which suggests an unusual genome sequencing; phylogenomic analyses tendency for ecological transitions (Ghobad- suggest that it is in a clade that also includes Nejhad et al. 2010; Lawrey et al. 2008). Ecology Gloeophyllales and Jaapiales (Fig. 14.1). A can sometimes be used to characterize genera, recent attempt to produce an infraordinal clas- but most clades in this order have mixtures of sification of Corticiales is based on a molecular nutritional modes. Entirely saprotrophic phylogeny using nuc-lsu rRNA sequences genera include Corticium, Giulia, and , (Ghobad-Nejhad et al. 2010). This analysis but Erythricium, which is mostly saprotrophic, resolved three groups that were recognized at also includes the plant pathogen Erythricium the family level using the existing names, salmonicolor. Vuilleminiaceae, Punctulariaceae, and Corticia- Most species of Limonomyces, Laetisaria, ceae. Until more sequences become available and are plant pathogens or endophytes and more specimens sequenced, this represents but are able to persist in the field as saprotrophs the best current hypothesis for a family (Andjic et al. 2005; Burdsall 1979; Burdsall et al. classification of the order. 1980; Stalpers and Loerakker 1982). However, Laetisaria also includes the lichen parasite L. Vuilleminiaceae Maire ex Lotsy: this clade contains lichenicola (Diederich et al. 2011). Marchandio- saprotrophic species that develop dendrohyphidia and myces, originally described for lichen parasites produce clamps and generally allantoid spores and (Diederich 1990; Etayo and Diederich 1996), is gelatinous fruiting bodies. It includes the genera Vuil- leminia and and a new genus, Australovuille- now known to include saprotrophic and folii- minia (for Vuilleminia coccinea). Based on their colous species (Diederich and Lawrey 2007; molecular phylogeny and incompatibility crossing Lawrey et al. 2007, 2008). There is also one tests, Ghobad-Nejhad et al. (2010) found that the so- lichen-forming species, Marchandiomphalina called core Vuilleminia species (. macrospora, V. foliacea (Lawrey et al. 2008; Palice et al. 2005), pseudocystidia, V. alni, V. comedens, V. megalospora) form a monophyletic group. These species are all dec- which seems to be most closely related to the orticating, produce a gelatinous fruiting body, and lichen parasite Marchandiobasidium aurantia- exhibit a unique 13 bp insertion in the ITS2. Other cum and the saprotrophic Erythricium laetum. described Vuilleminia species (V. cystidiata and V. Lichen mutualisms have evolved independently macrospora) were recovered in the Vuilleminia clade in Basidiomycota at least five times (Diederich but outside of the core Vuilleminia clade. Other species were recovered outside of the Vuilleminia clade and and Lawrey 2007; Diederich et al. 2003, 2011; reassigned to new genera, including V.(Punctulariop- Ertz et al. 2008; Fischer et al. 2007; Hodkinson sis) obduscens and V. (Punctulariopsis) subglobispora, et al. 2012; Lawrey et al. 2007, 2008, 2009; which were recovered in the Punctularia clade, and Nelsen et al. 2007; Redhead et al. 2002). Vuilleminia (Marchandiopsis) quercina, which was Marchandiomphalina foliacea is the only recovered in the Corticium clade. Punctulariaceae Donk: the family introduced by described lichen species in Corticiales. It Donk (1964) was intended to separate Punctularia spe- forms a foliose thallus structure and asexual cies from other corticioid fungi in the Aphyllophorales, goniocysts resembling soredia, but no sexual but few classifications recognized it, placing most cor- stages are known (Jørgensen 1989). ticioid species in the Corticiaeae. In the nuc-lsu rRNA Systematics: the monophyly of Corticiales phylogeny of Ghobad-Nejhad et al. (2010), this is a clade of saprotrophic species that produces clamps is strongly supported by molecular phyloge- and ellipsoid spores and includes the genera Punctu- nies, mostly based on rRNA gene sequences laria and and a new genus, Punctular- (Binder et al. 2005; Boidin et al. 1998; DePriest iopsis (for Vuilleminia subglobispora and Vuilleminia et al. 2005; Diederich et al. 2011; Ghobad- obducens). These species cause a vigorous white rot Nejhad et al. 2010; Hibbett et al. 2007; Langer compared to species in Corticium. 402 D.S. Hibbett et al.

Corticiaceae Herter: this family, originally con- been undersampled for sequences of Basidio- served against Vuilleminiaceae (Pouzar 1985) to repre- mycota, as well as for their fruiting bodies. sent a much broader circumscription than has emerged Systematics: the genus Jaapia Bres. was in recent molecular-based classifications, is now viewed by Ghobad-Nejhad et al. (2010) as a well-supported referred to the (Boletales) clade containing species with and without clamps, (Eriksson and Ryvarden 1976; Nannfeldt and including the type species of Corticium, and a variety Eriksson 1953) but later recognized as distinct of sexual and asexual genera with diverse nutritional from Boletales in rDNA analyses of Binder et al. modes (Erythricium, Galzinia, Giulia, Laetisaria, Limo- (2005) and Larsson (2007a). The latter study, nomyces, Marchandiobasidium, Marchandiomphalina, Marchandiomyces, Marchandiopsis, and Waitea). As based on analyses of sequences of 5.8S and nuc- mentioned by these authors, Corticiaceae is by far the lsu rRNA, confirmed that J. ochroleuca is a most diverse family in Corticiales, both morphologi- member of the same lineage as the type species, cally and ecologically. It also contains several polyphy- J. argillacea. A 6-gene phylogeny placed J. argil- letic genera in need of revision. The single most lacea as a sister group to Agaricomycetidae problematic clade, containing Marchandiomyces, Marchandiopsis, Limonomyces, and Laetisaria, is also (Atheliales, Boletales, Amylocorticiales, and the most interesting ecologically. Improving the inter- Agaricales) (Binder et al. 2010), but phyloge- nal resolution in this clade will not only resolve the nomic analyses place Jaapiales in a well- generic taxonomy of these groups but also help to supported clade with Corticiales and Gloeo- clarify some of the most interesting ecological transi- phyllales (Fig. 14.1). tions in Agaricomycetes.

K. Gloeophyllales J. Jaapiales Overview: Gloeophyllales Thorn is an odd Overview: Jaapiales Manfr. Binder, K.H. Larss. taxon with no morphological or ecological & Hibbett (Binder et al. 2010) is the smallest characters that unite the 6 genera and perhaps order of Agaricomycetes, with a single genus of 40 species (Hibbett et al. 2007; Kirk et al. 2008). just two species, Jaapia argillacea and J. ochro- The type genus, Gloeophyllum, is a bracket fun- leuca. Fruiting bodies of both species are resu- gus with resupinate, effused-reflexed, or pileate pinate, at first patchy, then thinly effused and fruiting bodies and poroid, daedaleoid, or monomitic, with thin-walled hyphae and fre- lamellate hymenophores (Gilbertson and quent clamp connections. Basidiospores are Ryvarden 1986). Boreostereum, Chaetodermella, narrowly fusoid (boletinoid) and cyanophilous. and Veluticeps (including Columnocystis)are Ecological diversity: both species of Jaapia corticioid to stereoid, with resupinate to fruit on wet, rotting wood on the margins of effused-reflexed fruiting bodies having a smooth lakes and streams and are collected infre- or rugose-wrinkled hymenophore (Chamuris quently (Eriksson and Ryvarden 1976). Only 1988; Eriksson and Ryvarden 1973; Nakasone J. argillacea is known in culture, and it is 1990b)(Fig.14.7b). Neolentinus and unreactive in tests for laccase, peroxidases, or produce agaricoid fruiting bodies with a central tyrosinase (Stalpers 1978). Thus, Jaapia species or eccentric , a convex to upturned , might be brown-rot saprotrophs or ECM, but and adnexed to decurrent lamellae (Redhead the biology of this group is unknown. BLAST and Ginns 1985)(Fig.14.7a). Most taxa are searches using the sequences of J. argillacea or dimitic, but some are monomitic, and others J. ochroleuca yield very few matches among trimitic. Simple clamp connections are constant environmental sequences, a rarity in the Agar- (most taxa), rare (some Veluticeps in culture), or icomycetes. At present, the only sequence absent (Boreostereum). The context is pallid in matches are to a few ITS sequences of uncul- Heliocybe and Neolentinus but brown in most tured fungi from permafrost. It is likely that other taxa and browning in KOH (turning green woody substrates in aquatic habitats have with KOH in Boreostereum) (Hibbett et al. 2007). Agaricomycetes 403

Sexuality ranges from homothallic (Boreoster- myces and Pileodon, which are segregates of eum, several Veluticeps) to heterothallic and Veluticeps (Nakasone 1990b), and Mycothele bipolar (Gloeophyllum, Heliocybe, Neolentinus) have been referred here, but no sequence data or tetrapolar (V. berkeleyi) (Ginns and Lefebvre are available. , from which 1993). Mycothele was segregated, has also been sug- Ecological diversity: members of Gloeo- gested as belonging in Gloeophyllales, but ana- phyllales are wood-decay fungi mostly causing lyses of rRNA gene sequences place it in the a brown rot of conifers, occasionally angios- core polyporoid clade of Polyporales (Kru¨ger perms, and frequently in wood in service. - and Gargas 2004). lentinus lepideus is known as the “train- Garcia-Sandoval et al. (2011) presented a 6- wrecker” for its propensity to decay wooden gene phylogenetic analysis of 18 species repre- railway trestles in bygone days (Redhead and senting the 6 genera accepted in Gloeophyllales. Ginns 1985), and species of Gloeophyllum com- Their results suggest that Gloeophyllum con- monly decay outdoor wooden structures such sists of at least two clades, one containing the as decks, playground equipment, and picnic type species, G. sepiarium, as well as G. stria- tables, and sometimes wooden joists and tim- tum, G. subferrugineum, and G. trabeum (all bers in homes (Gilbertson and Ryvarden 1986). species known from wood in service), and the Chaetodermella and Veluticeps cause brown other containing the type species of Osmoporus, rots of conifers, Heliocybe on angiosperms, O. odoratus, as well as Osmoporus protractus often in quite dry situations such as fence (both on exposed conifer wood in boreal- posts or rails and exposed, decorticated logs. subarctic environments) (Garcia-Sandoval However, Boreostereum, which seems to be a et al. 2011). Gloeophyllum mexicanum and sister group to the remainder of the order in a Gloeophyllum carbonarium were basal to 6-gene phylogenetic analysis (Garcia-Sandoval Osmoporus, and each might represent segregate et al. 2011), is associated with white rot of fire- genera upon further study. In addition, the type charred coniferous or angiosperm wood, species of Veluticeps (V. berkeleyi), Columno- although spot tests for laccases, peroxidases, cystis (C. abietina), and Chaetodermella (C. and tyrosinase in culture have been equivocal luna) formed a weakly supported clade for (Chamuris 1988; Nakasone 1990a). which the oldest generic name is Veluticeps.A Systematics: the core of Gloeophyllales was sequence of V. fimbriata was placed on a long recognized by Kim and Jung (2000) as Chaeto- branch that split the genus Neolentinus, but dermataceae, and the link between Gloeophyl- other analyses placed it together with the lum, Heliocybe, and Neolentinus was made by remaining species of Veluticeps, and Heliocybe Thorn et al. (2000). Studies by Binder et al. was recovered as sister to Neolentinus, leaving (2005) strongly supported early suggestions the possibility of its synonymy with Neolenti- (Hibbett and Donoghue 1995; Hibbett et al. nus undecided (Garcia-Sandoval et al. 2011). 1997b) that Gloeophyllum was set apart from Thus, for the moment we advocate recognition the true polypores and formed the basis for of Boreostereum, Gloeophyllum, Heliocybe, describing the Gloeophyllales (Hibbett et al. Neolentinus, Osmoporus, and Veluticeps in 2007). The polypore , which causes Gloeophyllales. a white rot of conifer wood in service, was included in the order when it was first described (Hibbett et al. 2007) on the basis of L. Russulales its clustering in the Gloeophyllum clade in ana- lyses of nuc-lsu rRNA sequence data by Kim Overview: Russulales currently includes more and Jung (2000, 2001), but it can now be than 1,700 described species (Kirk et al. 2008). excluded as a member of the core polyporoid This high number corresponds to an equally clade (Garcia-Sandoval et al. 2011). In addition astonishing diversity of fruiting body morphol- to the six genera known to belong to the order ogies (Fig. 14.8) and life strategies. From a phy- on the basis of molecular studies, Campylo- logenetic perspective, the dominant life form is 404 D.S. Hibbett et al. the skinlike, effused, and resupinate basidioma, ted. It can be highly intense, for example, by often developing out of sight at the underside of species in Stereum and , and decaying wood on the ground (Larsson and sometimes characteristic like the white pocket Larsson 2003). Examples include , rot produced by and ,andBoidinia. From ancestors spp. (Otjen and Blanchette 1984). Some species with such inconspicuous basidiomata, elaborate are capable of infecting living trees and perform fruiting structures have developed, for example, decay in roots or heartwood. Heterobasidion coralloid in , ,andArto- annosum is considered the most severe forest myces (Fig. 14.8c); reflexed and bracketlike as in pathogen in conifer forests in the Northern , Stereum, Laurilia (Fig. 14.8b), Hemisphere, causing economic losses of $1 bil- and (some species of) ;pileate- lion annually in the USA alone (Woodward stipitate mushrooms as in Russula, Lactarius et al. 1998). Other species with a potentially (Fig. 14.8a), and ; or sequestrate as pathogenic behavior in managed forests are in Macowanites (Fig. 14.8d, e)andLeucophleps. , Hericium erina- The hymenophore is most often smooth (e.g., ceus, Scytinostroma galactinum, and Stereum Peniophora, Stereum) or hydnoid (e.g., Auriscal- sanguinolentum. pium, Hericium), while a poroid hymenophore Another threat to is caused by configuration is comparatively rare (e.g., Alba- wood wasps from the family Siricidae living in trellus, Heterobasidion, ). A lamel- symbiosis with members of Amylostereum late hymenophore is known from (Slippers et al. 2003). The wasp female transfers (Lentinellus) and conidia of the fungus when she places eggs only. Many species have basidiospores with an inside the wood of stressed trees using her amyloid reaction of the spore wall, and for most needlelike ovipositor organ. Larvae then feed of them the amyloidity is combined with an on the fungus while mining through fungus- ornamented outline of the wall. infested wood. Infection by Amylostereum are- There is no obvious morphological synapo- olatum and its vector, Sirex noctilio, normally morphy for Russulales, but the presence of does not cause much damage within its native gloeoplerous hyphae or gloeocystidia with con- range in Europe and Asia. However, when the tents rich in sesquiterpenes has not been wasp was accidentally introduced into the demonstrated in any other basidiomycete Southern Hemisphere and in North America, order (Larsson and Larsson 2003). In a few infections in both exotic pine plantations and cases such structures seem to have been sec- native American pine stands have become ondarily lost (e.g., in ) or trans- severe (Nielsen et al. 2009; Slippers et al. 2001). formed into homologous structures such as ECM associations have developed indepen- the lactiferous hyphae in Stereum. The term dently within two Russulales lineages, in Rus- gloeocystidium refers to enclosed, bladderlike sulaceae and in . Molecular structures in fruiting bodies (Cle´menc¸on phylogenies suggest that in Russulaceae the 2004). Structures termed gloeocystidia have evolution proceeded from a saprotrophic to a been reported in many orders, but the specific mycorrhizal nutritional strategy and coincides type present in Russulales is associated with with the development of erect fruiting bodies unique vesicles with tubular invaginations, from effused ancestors (Larsson and Larsson which may constitute a synapomorphy for the 2003). Another trend confined to the same order (McLaughlin et al. 2008). In Russulales families is gasteromycetization (Albee-Scott the gloeoplerous hyphae and gloeocystidia have 2007), which involves the evolution of closed been suggested to serve as a chemical defense fruiting bodies adapted for a dryer climate and system against mycophagy (Sterner et al. 1985). animal dispersal. It is likely that change in life- Ecological diversity: the dominant nutri- style has driven the development of erect and tional strategy in Russulales is saprotophic closed fruiting structures since both are better decay of organic matter, primarily wood. In adapted for a soil-oriented life than the effused Russulales only white rot has been documen- structure typical for wood decayers. Agaricomycetes 405

Systematics: the potential presence of a has repeatedly taken place within Lactarius and Rus- russuloid lineage with the wide circumscription sula, respectively (Lebel and Tonkin 2007; Miller et al. accepted here was first discussed by Donk 2001; Nuytinck et al. 2004). Basal lineages are com- posed of saprotrophic taxa with corticioid fruiting bod- (1971). Oberwinkler (1977) elaborated on ies (, Gloeopeniophorella) (Larsson and Donk’s ideas and used Russulales as an exam- Larsson 2003). All species in Russulaceae have orna- ple of a higher-order group that contained mul- mented spores. tiple fruiting body morphologies. Molecular Albatrellaceae (six genera) is the second family data have confirmed that Russulales sensu where a mycorrhizal life strategy predominates. Species are stipitate-poroid (Albatrellus, ), Oberwinkler is a monophyletic group (Bruns effused-poroid (Byssoporia), or sequestrate (e.g., Myco- et al. 1998; Hibbett et al. 1997b, 2000; Larsson levis). Smith et al. (2013) present a phylogeny for the 2007b; Larsson et al. 2004; Binder et al. 2005). family. The most comprehensive phylogenies for Rus- (three genera) species have coralloid sulales have been published by Larsson and or effused that are mostly strongly hyd- noid. (lion’s mane, monkey’s head) Larsson (2003) and Miller et al. (2006). Larsson has been much used in Chinese folk medicine, and and Larsson (2003) identified ten well- modern studies have demonstrated the presence of supported lineages that can be understood as many medically active substances in this and related representing families; the main lineages are species (Lindequist et al. 2005; Mizuno 1999). A phy- briefly discussed in what follows. For many logeny for the genus is presented by Hallenberg et al. (2013). genera relationships are still not resolved. Auriscalpiaceae (four genera) includes lamellate (Lentinellus), stipitate-hydnoid (), and effused-hydnoid basidiocarps (Dentipratulum, Gloio- (14 genera) is dominated by corticioid don). forms with a smooth hymenophore and smooth basi- (four genera) is a small family of diospores. Most species fruit in exposed places like hard and robust wood-decaying species with either a living or recently dead trunks and branches and have poroid (, Heterobasidion) or a hydnoid morphological adaptations for resisting drought. hymenophore (Echinodontium). Examples include Stereum, sensu lato, : the clavarioid genus Artomyces is Aleurocystidiellum, and Xylobolus. Only Aleurodiscus sometimes included in Auriscalpiaceae, but that has been the subject of a detailed phylogenetic study arrangement is not unambiguously supported by (Wu et al. 2001). molecular data. Amylostereum could be placed in Bon- (16 genera) includes mainly corti- darzewiaceae but also recognized as a separate family cioid species with a smooth hymenophore and smooth (Binder et al. 2005). spores, not always with an amyloid reaction of the spore wall. Some genera are characterized by branched, dextrinoid skeletal hyphae (e.g., Scytinostroma, Var- aria). These genera are often referred to a separate family , but molecular data do not support its recognition. Also in this family, many spe- M. Agaricomycetidae cies grow rather exposed and decay dead but still attached branches (Peniophora, Scytinostroma). Agaricomycetidae contains two large and well- Closely related to Peniophora is Entomocorticium, known as symbionts of bark of the genus known orders, Agaricales and Boletales, as well Dendroctonus that cause great damage to pine forests as three small groups containing mostly corti- (Harrington 2005). Morphological studies of single cioid forms, Atheliales, Amylocorticiales, and genera are available, but a comprehensive phylogeny Lepidostromatales. Russulales seems to be the for the family is lacking. sister group of Agaricomycetidae. The clade Russulaceae (six genera) is the most species-rich family due to the high diversity seen in the mushroom containing Agaricomycetidae and Russulales genera Lactarius and Russula. Recent phylogenetic largely corresponds to Agaricales sensu Singer studies have shown that Russula is monophyletic, but (1986), which included four suborders, Agari- Lactarius in a traditional sense is not (Buyck et al. cineae, , Russulineae, and Polypori- 2008). The latter group is now divided into Lactarius, neae. The latter included agaricoid forms in Lactifluus, and . The many genera with sequestrate species recognized earlier (e.g., Arcange- Lentinus and other genera that are now liella, Macowanites) are now considered examples of known to be distributed among Polyporales, adaptations to a dry habitat and animal dispersal that Gloeophyllales, and Agaricales. 406 D.S. Hibbett et al.

1. Atheliales and Lepidostromatales dae incertae sedis based on mixed support values from rDNA analyses (Ertz et al. 2008). Overview: Atheliales Ju¨lich in a broad sense In a recent multigene study this family was includes 22 genera with 110 described species recognized as the order Lepidostromatales, (Kirk et al. 2008). However, several genera of including the genera Lepidostroma, Sulzbacher- Atheliales are polyphyletic (e.g., some Athelia, omyces, and Ertzia (Hodkinson et al. 2013). A Athelopsis, and species) with taxonomic revision of on the species in Agaricales, Amylocorticiales, generic level is needed because new taxa are Cantharellales, and Polyporales (Binder et al. being described (Kotiranta et al. 2011) and pre- 2005, 2010; Ertz et al. 2008; Larsson 2007b; viously unknown lineages are being added to Larsson et al. 2004; Matheny et al. 2006; Ober- the family (Sjo¨kvist et al. 2012). winkler 2012). Many athelioid species produce loosely connected resupinate fruiting bodies lacking conspicuous morphological differentia- 2. Amylocorticiales tion on various substrates, including branches, wooden debris, and mosses (Fig. 14.9d) (Lars- Overview: Amylocorticiales K.H. Larss., Manfr. son et al. 2004). vitellina forming Binder & Hibbett is a recently described order stipitate-stereoid basidiocarps was recently (Binder et al. 2010)thatincludesroughly70 separated from Cantharellales and placed in species. Taxonomic concepts at the generic Atheliales (Sjo¨kvist et al. 2012). The lichenized level are still in flux (Binder et al. 2005, 2010; Lepidostromataceae was originally placed in a Buyck et al. 2012;Gorjo´netal.2011; Larsson sister-group relationship to Atheliales and 2007b; Niemela¨ et al. 2007;Zmitrovichand included three species that produce clavarioid Spirin 2002). Species of the nine genera accepted basidiocarps similar to Multiclavula spp. in in Amylocorticiales usually form corticioid and Cantharellales (Ertz et al. 2008). resupinate fruiting bodies and produce smooth, Ecological diversity: Atheliales is not a merulioid, or sometimes poroid hymenophores species-rich group, but it is pervasive in terres- (Amylocorticiellum, , Anomo- trial ecosystems. Some Athelia species parasit- loma, Anomoporia, , Serpulo- ize cyanobacteria, green algae, and myces)(Fig.14.9b). Others have evolved more (Oberwinkler 1970; Yurchenko and Golubkov elaborate fruiting bodies, including multistoried 2003), and it has been suggested that the pileate-stipitate structures (, lifestyle of the lichen-forming Lepidostromata- the pagoda fungus), pendant fan-shaped fruiting ceae can be considered a similar form of inter- bodies with wrinkled gill-like hymenophores action (Oberwinkler 2012). Other Athelia, (Plicaturopsis crispa), or hydnoid hymeno- Athelopsis, and Tretomyces spp. produce white phores ( pendulus). Anatomical char- rot on various trees, debris, leaf litter, grasses, acters in Amylocorticiales are also diverse. The and ferns (Eriksson and Ryvarden 1973; Kotir- basidiospores are either thin- or thick-walled, anta and Saarenoksa 2005; Kotiranta et al. smooth, ellipsoid, cylindrical, or allantoid, and 2011). Brown rot is absent in Atheliales (Binder most react positively (amyloid) to Melzer’s et al. 2010). The Athelia anamorph Fibulorhi- reagent. All hyphal systems are monomitic (i.e., zoctonia forms symbioses with by pro- consist of generative hyphae only) and nodose ducing sclerotia that mimic termite eggs septate; however, this character combination is (Matsuura et al. 2000). , Byssocorti- not synapomorphic for Amylocorticiales and cium, Piloderma, and spp. are ECM occurs in other groups (e.g., Polyporales). Cysti- with Pinaceae and Fagaceae and are often dom- dia are rare in Amylocorticiales. inant in ECM fungal communities (Erland and Ecological diversity: species placed in Taylor 1999; Lilleskov et al. 2004). Amylocorticiales are predominantly sapro- Systematics: Atheliales currently includes trophic or, rarely, biotrophic. The modes of Atheliaceae as a single family. Lepidostromata- wood decay include brown rot (e.g., Amylocor- ceae had been formally left in Agaricomyceti- ticium, Anomoporia, Podoserpula) and white Agaricomycetes 407 rot (e.g., , Irpicodon, Plicaturopsis). with gilled hymenophores, such as or molle (Leucogyrophana mol- , have evolved at least five times lis) is of economic importance as a causal agent from resupinate ancestors (Binder et al. 2005, of brown rot in timber (Mattsson et al. 2010; 2010). taxi is the only species Niemela¨ et al. 2007). The decay strategy of Ser- developing polyporelike basidiocarps (Larsson pulomyces has not been studied in detail. ECM 2007b), but no coralloid or clavarioid forms are forms are seemingly absent in Amylocorti- known in the order. The fruiting bodies of ciales, but it has been suggested that Anomo- Boletales are specifically attacked by the asco- loma flavissimum and Podoserpula pusio may mycete anamorph genus Sepedonium (teleo- represent transitions to ECM symbioses morph Hypomyces), suggesting some degree (Bougher and Syme 1998; Niemela¨ et al. 2007). of coevolution between parasites and hosts “Athelia” rolfsii (anamorph rolfsii) (Douhan and Rizzo 2003; Sahr et al. 1999). is a serious soilborne pathogen, also known as The morphological characters of Boletales Southern blight, that infects more than 500 have been studied intensively (Agerer 1999; plant species, including , potato, and Arpin and Ku¨hner 1977; Both 1993; Corner (Punja 1985). 1972; Horak 2004, 2011; Moser 1983; Pegler Systematics: Amylocorticiaceae Ju¨lich is and Young 1981; Singer 1986; Smith and Thiers the single family in Amylocorticiales to date, 1971; Watling 1970), but there is no synapo- and a major taxonomic revision on the generic morphic trait for the order as a whole. Boletales level is needed. Amyloathelia crassiuscula, species produce unique pigments and colorless allantosporum, Anomoporia compounds during secondary metabolism, and kamtschatica, Athelia rolfsii, Athelopsis lacer- the terphenyl quinone atromentin plays an ata, Leptosporomyces septentrionalis, and essential role as building block for the synthesis Hypochniciellum molle are distinct lineages in of derivatives, including pulvinic acids (e.g., Amylocorticiales, but they do not represent the variegatic acid and xerocomic acid), cyclopen- generic types. tenones, grevillins, and other substances (Besl and Bresinsky 1977, 1997; Besl et al. 1986; Bresinsky 1974; Bresinsky and Orendi 1970; 3. Boletales Gill and Steglich 1987). An atromentin pathway has evolved independently in Thelephorales, Overview: Boletales E.-J. Gilbert is one of the but it produces only simple terphenyl larger orders of fleshy Agaricomycetes, includ- quinones, not the structurally more complex ing 17 families, 88 genera, and roughly 1,400 pigments (Besl and Bresinsky 1997). species (Binder and Hibbett 2006; Kirk et al. Ecological diversity: members of Boletales 2008). The typical fruiting body of a is have a worldwide distribution in forest ecosys- pileate-stipitate with a tubular (e.g., Boletus, tems, with biodiversity hot spots in Southeast ) (Fig. 14.9e) or sometimes gilled hyme- Asia and North America (Corner 1972; Singer nophore (Paxillus, ). Gasteroid 1965; Smith and Thiers 1971). The major nutri- forms (, , ) tional modes of Boletales include brown-rot have evolved several times independently saprotrophy, ECM symbioses, and mycopara- from this morphology (Binder and Bresinsky sitism; biotrophic plant pathogens and white- 2002; Binder and Hibbett 2006; Bruns et al. rot fungi are not known (Binder and Hibbett 1989; Thiers 1984; Wilson et al. 2011). Roughly 2006). It has been suggested that brown rot is 77 % of the described species produce pileate- the ancestral lifestyle of Boletales, having a sin- stipitate fruiting bodies (Binder et al. 2010). In gle evolutionary origin in the early branching addition, there are resupinate forms with lineages (Binder and Hibbett 2006). Based on smooth or warted (, Serpula) the unique capability of brown-rot-producing (Fig. 14.9f), merulioid (Leucogyrophana), and Boletales to selectively depolymerize micro- toothed () hymenophores, and it crystalline cellulose, which weakens the has been suggested that pileate-stipitate fungi strength of wood, this form of wood decay has 408 D.S. Hibbett et al. also been called Coniophoraceae-type rot ectomycorrhizae of the closely related Suillus (Ka¨mmerer et al. 1985; Nilsson and Ginns and Rhizopogon by penetrating their rhizo- 1979) to separate it from other brown-rot morphs (Agerer 1990, 1999; Miller 1964; Olsson types. Most brown-rot-causing species contrib- et al. 2000), thereby circumventing competition ute to carbon sequestration in conifer forests for host plants (Binder and Hibbett 2006). Pseu- (Tapinella, Pseudomerulius), but a few have doboletus parasiticus in produces its specialized on human-built timber environ- fruiting bodies on (a gas- ments. The so-called cellar fungus Coniophora teroid bolete) while eroding the gleba of the puteana and especially the so-called dry rot host (Binder and Hibbett 2006). Other putative fungus cause significant mycoparasites from the basal lineages of Bole- damage in wooden building structures taceae include the sister taxa (Schmidt and Kebernik 1989; Schmidt et al. and (Nuhn et al. 2013). 2002), and the ecological diversification and Systematics: Boletales includes five subor- structure of geographical lineages of these ders that have been described based on dispa- aggressive decayers have been studied in detail rate characteristics and methods: Boletineae, (Eastwood et al. 2011; Kauserud et al. 2007a, b; Suillineae, , Tapinellineae, Skrede et al. 2011; Watkinson and Eastwood and Coniophorineae (Binder and Hibbett 2012). is also a prime example of 2006). Boletineae was first introduced by transitions from brown-rot to ectomycorrhiza Gilbert (1931) using fruiting body morphology associated with major morphological changes. and spore shape as distinctive characteristics. species (pileate-stipitate fruiting This suborder included all species with tubular bodies with gilled hymenophores) and Gymno- hymenophores at that time and a few species paxillus species (gasteroid) are derived from with gilled hymenophores. Suillineae was sepa- within Serpula and form ectomycorrhizae with rated later from Boletineae based on unique Eucalyptus and (Bresinsky et al. pigments occurring in this group (Besl and 1999; Jarosch 2001; Skrede et al. 2011). Bresinsky 1997). In addition, numerous resupi- Approximately 90 % of species in Boletales nate and paxilloid taxa producing a brown rot are involved in ECM symbioses, particularly were known to be closely related to Boletales with Betulaceae, Caesalpiniaceae, Dipterocarpa- based on their pigments (Besl et al. 1986), but ceae, Fagaceae, Myrtaceae, Nothofagaceae, Pina- they remained incertae sedis. The morphology ceae, and Salicaceae, or in arbutoid of belowground rhizomorphs helped to for- with Ericaceae (Newman and Reddell 1987; mally place these taxa in Tapinellineae and Rinaldi et al. 2008; Tedersoo et al. 2010). Boletus Coniophorineae (Agerer 1999), which was sup- and spp. show an increased tendency ported by early major studies using DNA to associate with specific hosts; for example, sequences (Bruns et al. 1998; Kretzer and forms ectomycorrhizae with Bruns 1999). Sclerodermatineae was described Betula (Singer 1967). Suillus, , based on nuc-lsu rRNA sequences (Binder and ,andRhizopogon spp. in the Bresinsky 2002) integrating the gasteroid Scler- suborder Suillineae are almost exclusively asso- odermatales into Boletales. ciated with Pinaceae, which is probably the old- Resupinate taxa are still a source of taxo- est clade of ECM partners for Boletales (Hibbett nomic uncertainty in Tapinellineae and Con- and Matheny 2009). Most ECM species are iophorineae, particularly concerning the placed among the Boletaceae (roughly 400 plus polyphyletic genus Leucogyrophana (Binder species), which include highly prized edibles et al. 2010; Jarosch and Besl 2001). Conio- such as (porcini). phorineae, including three clades of Leucogyr- Mycoparasites in Boletales represent tran- ophana, has been resolved as a monophyletic sitions from the ECM lifestyle (Binder and group, but without significant statistical sup- Hibbett 2006) and have evolved at least twice port (Binder et al. 2010). Together, Boletineae, independently. Gomphidius and Chroogomphus Sclerodermatineae, and Suillineae form the spp. are capable of parasitizing the established largest clade, including the overwhelming Agaricomycetes 409 majority of ECM taxa (except Austropaxillus the arrangement of the covering layers of the and ) and mycoparasites. Hyd- fruit body (pileipellis, stipitipellis) and the nomerulius pinastri (formerly Leucogyro- hymenophoral trama; and the presence of phana), a single species that is not placed in specialized structures (cystidia, setae) (Cle´men- any of the suborders, is sister to the remain- c¸on 2004; Reijnders and Stalpers 1992; Singer ing Boletineae members (Jarosch and Besl 1986). All these characters have played a central 2001). Current research is focused on the role in defining the approximately 400 genera ecology of Sclerodermatineae (Wilson et al. and 30 families in the order, but much of the 2007, 2012) and the taxonomic structure of taxonomy of the order is currently in flux as Boletineae, especially the Boletaceae. Mono- data from molecular phylogenies become graphic work has led to a better definition incorporated. A promising pool of micromor- of Boletus and its being restricted to the B. phological features that may be useful as future edulis group (Dentinger et al. 2010; Nuhn systematic markers in Agaricales is the complex et al. 2013), the revision of gilled in of characters related to conidiogenesis in ana- Phylloporus (Neves et al. 2012), and Xeroco- morphic stages (Walther et al. 2005). Since mus, which has been split into several new monosporic cultures obtained from basidios- genera (Sˇutara 2008). Since 2007, 14 new pores are usually needed to study these char- genera have been described in Boletaceae acters, this complex of features has clearly been (Desjardin et al. 2008, 2009; Halling et al. understudied. 2007, 2012a, b; Hosen et al. 2013; Lebel et al. The second major morphological compo- 2012; Li et al. 2011; Orihara et al. 2010;Sˇutara nent of Agaricales are the secotioid and gaster- 2008; Trappe et al. 2013; Zeng et al. 2012). oid forms, including false truffles, puffballs, and bird-nest fungi, that have evolved repeat- 4. Agaricales edly in different lineages within the order (e.g., Lycoperdon) (Fig. 14.9c). Additional morphol- Overview: Agaricales (Underwood 1899) ogies that can be found in the order include includes over 13,000 described species (Kirk resupinate (e.g., ), coralloid et al. 2008), making it the largest order of (e.g., Clavaria), cyphelloid (e.g., Henningso- Agaricomycetes. Despite being one of the myces), pileate with poroid (e.g., ), most conspicuous and comparatively better or tubular hymenophores (e.g., Fistulina). studied groups of fungi, an immense amount There is no morphological synapomorphy that of the actual diversity remains undescribed, unites the Agaricales, and the typical pileate- hiding under commonly used names that stipitate gilled mushroom morphology that molecular data have revealed to be clusters dominates the order also occurs in other orders of morphologically cryptic species (e.g., Ama- of Agaricomycetes. nita muscaria) (Geml et al. 2006) and part Ecological diversity: two ecological roles of hyperdiverse lineages with over 2,000 characterize the majority of Agaricales species: estimated species such as (e.g., saprotrophy and ECM symbiosis. Saprotrophs Harrower et al. 2011). can be broadly subdivided into soil/litter/dung Agaricales is dominated by pileate-stipitate fungi (e.g., Agaricus, Coprinopsis, ) forms with lamellate hymenophores (e.g., Ama- and wood decayers (e.g., , Pleurotus), nita, Agaricus, s.l., , Lepiota, but the exact roles and capabilities of members ) (Fig. 14.9a), but there is wide vari- of both ecological guilds remain largely ation on this basic fruit-body morphology unknown, although the emerging field of fungal regarding characters such as the size of the genomics is bringing new insights into these , the presence of veils (universal aspects (e.g., Morin et al. 2012). There have and partial), gill attachment, and spore-print been at least ten independent and asynchro- color (white, brown, purple-brown, black, nous origins of the ECM symbiosis in Agari- pink). Microscopically, there is also a great cales, involving associations with a great variety diversity of characters, including spore size, of vascular plants (Matheny et al. 2006; Ryberg shape, ornamentation, and chemical reactions; and Matheny 2012; Tedersoo et al. 2012). There 410 D.S. Hibbett et al. are many additional ecological roles in Agari- graphs are lacking for all major genera of Agar- cales, including plant pathogens (e.g., Moni- icales. liophthora/) (Meinhardt et al. 2008), mycoparasites (e.g., ) (Matheny Agaricoid clade: this clade is well supported in the study and Griffith 2010), basidiolichens (e.g., Liche- of Matheny et al. (2006) and is dominated by gilled pileate-stipitate forms with pigmented spores (brown, nomphalia and Dictyonema) (Dal-Forno et al. purple-brown, black). Exceptions to this general pattern 2013; Lawrey et al. 2009), attine ant cultivars include (1) white-spored taxa in (Lepiota (/ clade) (Mueller and allied genera), Cystodermateae, and et al. 2005), or termite cultivars (Termitomyces) (e.g., Laccaria); (2) secotioid and gasteroid taxa that (Aanen et al. 2002). The ecology of many Agar- have repeatedly evolved in different lineages (e.g., Agar- icaceae, Cortinarius); and (3) cyphelloid forms in the icales species remains poorly understood, and genera Pellidiscus and (Bodensteiner et al. new insights from molecular and isotopic data 2004). At least six ECM lineages are included in this are challenging long-standing views even in clade: Cortinarius, (and allied sequestrate relatively well-studied groups. For example, taxa), Inocybaceae, Laccaria (and allied sequestrate the so-called waxcaps ( sensu lato) taxa), , and some groups of Hymenogas- traceae (Alnicola, ). Most other members of have historically been considered saprotrophs the agaricoid clade are saprotrophs associated with the but now are thought to be involved in some litter layer and similar substrates (e.g., Agaricus, Copri- kind of biotrophic association (Lodge et al. nopsis) or wood decayers (e.g., ). Taxa with 2013; Seitzman et al. 2011). recent phylogenetic studies include Agaricaceae Systematics: three landmark papers pub- (Vellinga 2004; Vellinga et al. 2011), (To´ et al. 2013), (Frøslev et al. 2005;Garnica lished in recent years have redefined the taxo- et al. 2005;Peintneretal.2004), Crepidotaceae (Aime nomic organization of Agaricales species and et al. 2005), Cystodermateae (Saar et al. 2009), Gymno- their closest relatives: (1) Moncalvo et al. (2002) pileae (Guzma´-Da´valos et al. 2003), Hebelomateae presented the first broad phylogeny of the (Boyle et al. 2006;Moreauetal.2006), Inocybaceae order, based on nuc-lsu rRNA, which resolved (Matheny 2005;Rybergetal.2010), (Zhao et al. 2007), (Nagy et al. 2012), 117 clades and outlined conflicts with tradi- (Ramı´rez-Cruz et al. 2013), Tubarieae tional morphologically defined groups; (2) (Gulden et al. 2005), and several secotioid/gasteroid Matheny et al. (2006) presented the first major taxa usually nested within agaricoid relatives (Larsson multilocus overview of the order, including the and Jeppson 2008; Lebel and Syme 2012). protein-coding genes rpb1, rpb2, and tef1- Tricholomatoid clade: pileate-stipitate gilled mush- rooms with white or pink spores dominate this lineage, alpha, and defined six major infraordinal which includes four traditionally recognized families: clades: agaricoid, tricholomatoid, marasmioid, , , , and a hygrophoroid, pluteoid, and plicaturopsidoid restricted version of . A fifth lineage clades, although support for some of these includes the ECM and the saprotrophic groupings was weak; (3) Binder et al. (2010) . Additional ECM origins have also occurred in Entoloma, ,andTricholomata- built on the data set of Matheny et al. (2006) ceae. The clade also includes soil/litter saprotrophs (e.g., and formally recognized the plicaturopsidoid ), mycoparasites (e.g., Asterophora), wood clade as the order Amylocorticiales. decayers associated with white rot (e.g., )or Recent monographs on Agaricales include brown rot (e.g., ), and termite cultivars (Termi- the titles of the Fungi Europaei series on tomyces). Entolomataceae remains one of the most dis- tinct groups in Agaricales because of the pinkish spores Agaricus (Parra-Sa´nchez 2008), Amanita (Neville that are warted, ridged, or angular. Co-David et al. (2009) and Poumarat 2004), - recognize two broadly defined genera in the family (Ento- (Hausknecht 2009), and the family - loma and ), while other authors prefer to recog- ceae (Noordeloos 2011). Outside Europe mono- nize several narrowly defined genera (Baroni and graphic work at the continental scale is rare, a Matheny 2011; Largent et al. 2011). The Lyophyllaceae has also been reviewed using molecular phylogenies situation that may change with large-scale cata- (Hofstetter et al. 2002).Thetaxonomyofmanytradition- loging efforts that include the use of molecular ally recognized genera (e.g., Tricholoma, Clitocybe, data, such as the North American Mycoflora , Mycena) is still in flux. project (Bruns 2012). Modern global mono- Marasmioid clade: this lineage is dominated by white-spored saprotrophic species associated with Agaricomycetes 411 wood or leaf-litter substrates (e.g., Lentinula, Maras- diomycetes, with eight informally named mius, ), but important plant pathogens also clades, that was based on 25 published and belong here (e.g., Armillaria, ). Schizo- unpublished analyses (Hibbett and Thorn phyllaceae (Fistulina, Schizophyllum) and - ceae, dominated by cyphelloid forms, were recovered 2001). The present chapter cites nearly 300 as part of the marasmioid clade by Matheny et al. phylogenetic studies, many combining rRNA (2006) but as independent lineages in Binder et al. and protein-coding genes, and a handful of (2010). Resupinate genera (e.g., ) also phylogenomic analyses. Twenty strongly sup- occur in the marasmioid clade. Representative taxa ported, mutually exclusive clades of Agarico- with recent phylogenetic studies include a general over- view of marasmioid/gymnopoid fungi (Wilson and mycetes are recognized as orders. Numerous Desjardin 2005), (Kirchmair et al. 2004), studies, most of which are not cited here, have (Mata et al. 2004), , and Cri- addressed species- and genus-level relation- nipellis (Kerekes and Desjardin 2009; Wannathes et al. ships within these groups. Nevertheless, the 2009), and genera in the Xerula/ com- classification of Agaricomycetes is far from plex (Petersen and Hughes 2010). Hygrophoroid clade: in the analyses of Matheny complete. There are weakly supported nodes et al. (2006) this clade includes an expanded version of throughout the phylogeny, and the catalog of as traditionally defined plus the club described species is thought to be a tiny and coralloid fungi in the families and fraction of the actual diversity in the group . However, these two families were later (Blackwell 2011). resolved as a separate lineage in Agaricales (Binder et al. 2010). The family Hygrophoraceae was recently An overarching challenge of fungal system- extensively studied and redefined by Lodge et al. (2013) atics is to capture and integrate the massive and now includes 18 genera, including the ECM Hygro- volumes of data flowing from taxonomy, phy- phorus, several segregates from Hygrocybe sensu lato, logenetics, genomics, and molecular ecology. and a diverse clade of basidiolichens such as Dictyo- Unfortunately, some common practices make nema s. l. (Dal-Forno et al. 2013). All members of the family are now assumed to be involved in some kind of it difficult to combine the products of different biotrophic relation, but its exact nature remains areas of research. For example, curated obscure in most cases (Seitzman et al. 2011). sequence databases are important for the iden- Pluteoid clade: this clade received weak support in tification of environmental sequences (Ko˜ljalg Matheny et al. (2006), and its limits and composition et al. 2013), so it is unfortunate that many require further study. The grouping of the and is well supported in most phylogenies recent species descriptions have been published (Justo et al. 2011; Matheny et al. 2006). The aquatic without sequences (Hibbett et al. 2011). It is gasteromycete is also part of this core also unfortunate that only about 17 % of pub- pluteoid group in some analyses (Matheny et al. lished phylogenies, including those from fungal 2006). , , and the genus studies, are available in electronic form (not are, in some topologies, recovered as closely related to the core pluteoid genera but graphics files, but treefiles, in Newick or other not always with statistical support or in a consistent formats), which limits efforts to assemble max- position. Important taxonomic revisions include arti- imally inclusive phylogenies and combine them cles on the Pluteaceae (Justo et al. 2011), Melanoleuca with taxonomic hierarchies (Collins et al. 2013; (Sa´nchez-Garcı´a et al. 2013; Vizzini et al. 2012), Pleur- Drew et al. 2013). To approach a comprehen- otaceae (Thorn et al. 2000), and sequestrate forms in Amanita (Justo et al. 2009). The iconic genus Amanita sive phylogenetic classification of Agaricomy- has received considerable attention in relation to bio- cetes and other Fungi, it will be necessary to geography (Geml et al. 2006), invasive species (Pringle increase the pace of taxon discovery, encourage et al. 2009), and transitions from saprotrophic to ECM researchers to generate and deposit sequences, nutrition (Wolfe et al. 2012). alignments, trees, and associated metadata (Hyde et al. 2013), and create new bioinformat- ics tools to synthesize the prodigious output of the fungal systematics community. III. Conclusions Acknowledgements The authors are grateful to Sigis- fredo Garnica, Romina Gazis, Igor Grigoriev, Chris- The previous edition of The Mycota included a tiane Karasch-Wittmann, Urmas Ko˜ljalg, Francis preliminary phylogenetic outline of Homobasi- 412 D.S. Hibbett et al.

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