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fungal ecology xxx (2014) 1e10

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Short Communication How many fungi make sclerotia?

Matthew E. SMITHa,*, Terry W. HENKELb, Jeffrey A. ROLLINSa aUniversity of Florida, Department of , Gainesville, FL 32611-0680, USA bHumboldt State University of Florida, Department of Biological Sciences, Arcata, CA 95521, USA article info abstract

Article history: Most fungi produce some type of durable microscopic structure such as a that is Received 25 April 2014 important for dispersal and/or survival under adverse conditions, but many also Revision received 23 July 2014 produce dense aggregations of tissue called sclerotia. These structures help fungi to survive Accepted 28 July 2014 challenging conditions such as freezing, desiccation, microbial attack, or the absence of a Available online - host. During studies of hypogeous fungi we encountered morphologically distinct sclerotia Corresponding editor: in nature that were not linked with a known . These observations suggested that Dr. Jean Lodge many unrelated fungi with diverse trophic modes may form sclerotia, but that these structures have been overlooked. To identify the phylogenetic affiliations and trophic Keywords: modes of sclerotium-forming fungi, we conducted a literature review and sequenced DNA Chemical defense from fresh sclerotium collections. We found that sclerotium-forming fungi are ecologically Ectomycorrhizal diverse and phylogenetically dispersed among 85 genera in 20 orders of Dikarya, suggesting Plant pathogens that the ability to form sclerotia probably evolved 14 different times in fungi. Saprotrophic ª 2014 Elsevier Ltd and The British Mycological Society. All rights reserved. Sclerotium

Fungi are among the most diverse lineages of eukaryotes with features such as a hyphal thallus, non-flagellated cells, and an estimated 5.1 million species (Blackwell, 2011). They are the the production of (Stajich et al., 2009). However, principle saprotrophs in most terrestrial biomes and play because of their cryptic lifestyles within environments such important ecological and economic roles as plant pathogens as plants and soil, the ecology and evolutionary history of and mutualists. Fungi are found in all terrestrial ecosystems many fungi remains poorly understood. and they use a variety of strategies to colonize appropriate Almost all fungi produce some type of durable, quiescent substrata and survive unfavorable conditions (Blackwell, microscopic structure such as a spore that is important for 2011). They have significant impacts on the biology of plants dispersal and/or survival under adverse conditions (Stajich because they are the most economically significant plant et al., 2009). However, some fungi also produce dense aggre- pathogens, serve as mycorrhizal and endophytic symbionts, gations of fungal tissue called sclerotia (Willetts, 1971). These and act as key players in nutrient cycles (Schumann, 1991; persistent structures help fungi to survive challenging con- Rodriguez et al., 2009; Hobbie and Hogberg, 2012). Two fun- ditions such as freezing temperatures, desiccation, microbial gal phyla, and , comprise the attack, or the long-term absence of a host (Townsend and subkingdom Dikarya, a diverse group with ca. 100,000 Willetts, 1954; Coley-Smith and Cooke, 1971). Sclerotia are described species (James et al., 2006). Most Dikarya share key highly variable in their morphology (Fig 1). Some have a hard,

* Corresponding author. Tel.: þ1 352 273 2837; fax: þ1 352 392 6532. E-mail address: trufflesmith@ufl.edu (M.E. Smith). http://dx.doi.org/10.1016/j.funeco.2014.08.010 1754-5048/ª 2014 Elsevier Ltd and The British Mycological Society. All rights reserved.

Please cite this article in press as: Smith, et al., How many fungi make sclerotia?, Fungal Ecology (2014), http://dx.doi.org/ 10.1016/j.funeco.2014.08.010 2 M.E. Smith et al.

Fig 1 e Morphologically variable sclerotia found in soil, leaf litter, and decayed wood in natural forest habitats of North and South America: (A) Ceriporia sp. (MES 332; Polyporales) from decayed wood on the forest floor in Pigsah National Forest, North Carolina, USA; (B) Entoloma sp. (MES 347; ) from soil in a tropical rainforest dominated by leguminous ectomycorrhizal trees, Guyana; (C) Cheilymenia sp. (MES 313; ) from soil in mixed woods near Cherryfield, Maine, USA; (D) unknown species of Amylocorticiales (MCA 3949) from soil and leaf litter in a tropical rainforest in Guyana; (E) Boletus sp. (MES 260; ) from soil and leaf litter in angiosperm-dominated forest in Lexington, Massachusetts, USA. Identities of illustrated sclerotia were determined based on ribosomal DNA sequence comparisons with GenBank. Scale bars [ approximately 10 mm.

melanized rind enclosing compact, undifferentiated hyphae fungi such as sclerotiorum produce sexual fruiting while others lack a rind (Willetts, 1971). Some species make structures directly on sclerotia (Bolton et al., 2006) whereas round, determinate sclerotia but others have indeterminate others such as Pteromyces flavus (¼Aspergillus flavus) produce forms where the shape and size are influenced by resources fruiting bodies within sclerotia (Horn et al., 2009). Still others, and environmental conditions (Chet and Henis, 1975). Some such as Boletus rubropunctus, produce fruit bodies and sclerotia sclerotia are produced inside of host tissues; at different times or in different places (Smith and Pfister, produces sclerotia in grass florets after it has destroyed the 2009). plant cells (Douhan et al., 2008) and Although sclerotia have been documented in several fun- colonizes caterpillars and transforms their tissues into a gal lineages, sclerotium formation is primarily recognized as a sclerotium (Xing and Guo, 2008). In contrast, some fungi make key life history trait in several necrotrophic plant pathogens sclerotia that are spatially separated from hosts (e.g. Phyma- (e.g. Sclerotium rolfsii, solani, M. phaseolina, P. omni- totrichopsis omnivora forms sclerotia deep in soil e Lyda, 1984). vora, S. sclerotiorum). Collectively, these devastating host gen- Sclerotia also range in size from “microsclerotia” <1mm eralist pathogens are responsible for hundreds of millions of across, as in the plant pathogen Macrophominia phaseolina dollars in global crop losses annually (Aycock, 1966; Parmeter, (Short and Wyllie, 1978), to the massive sclerotia of Polyporus 1971; Purdy, 1979; Mulrean et al., 1984). For example, S. scle- mylittae that reach over 40 cm in diameter (Macfarlane et al., rotiorum and S. rolfsii each attack >400 plant species, including 1978). Sclerotia putatively serve a resource-storage and sur- major crops such as peanuts, potatoes, and soybeans, and can vival role in all sclerotium-forming fungi. However, some cause up to 100 % yield losses (Jenkins and Averre, 1986;

Please cite this article in press as: Smith, et al., How many fungi make sclerotia?, Fungal Ecology (2014), http://dx.doi.org/ 10.1016/j.funeco.2014.08.010 How many fungi make sclerotia? 3

Bowen et al., 1992; Cintas and Webster, 2001). For these and schematic phylogeny based on Hibbett et al. (2007) with phy- other sclerotium-forming pathogens, survival is tightly linked logenetic positions of new or revised orders inferred from with sclerotium formation so sclerotia eradication is critical LoBuglio and Pfister (2010), Schoch et al. (2009a, 2009b), Binder for disease control (Coley-Smith and Cooke, 1971). Fur- et al. (2010), Zhang et al. (2011), Toome et al. (2013), Boehm thermore, the ecology of these fungi cannot be fully under- et al. (2009), Campbell et al. (2009), and Padamsee et al. stood without understanding the biology of sclerotium (2012). All but two fungal species, salvinii and formation. Verticilium dahliae, were easily resolved at the ordinal level Although management of serious plant pathogens is an based on data from published references (Table 2) and Index important rationale for studying sclerotium formation, there Fungorum (www.indexfungorum.org/). are nevertheless several other compelling reasons. First, We documented reports of sclerotium formation in species many sclerotia lie dormant in soil, leaf litter, or wood for from 85 fungal genera in at least 20 orders of Basidiomycota months, so they must survive attacks from a wide variety of and Ascomycota (Table 2, Fig 2). Since only one representative natural enemies, including bacteria, other fungi, and inver- sclerotium-forming species from each was recorded, tebrates (Willetts, 1971; Papavizas, 1977; Matsumoto and we cannot accurately estimate the number of sclerotium- Tajimi, 1985). The mechanisms that allow sclerotia to sur- forming species. However, we observed that many genera vive in soil despite ongoing biotic assault are not well known, with one sclerotium-forming species also contain others. but evidence from well-studied species (e.g. S. sclerotiorum, C. Also, despite our limited sampling of sclerotia, we found a purpurea) suggests that most sclerotia contain biologically wide diversity of sclerotium-forming fungi in nature and active secondary metabolites (Morrall et al., 1978; Demain, documented at least three genera for which sclerotium for- 1999; Schardl et al., 2006; Ikewuchi and Ikewuchi, 2008; mation had not previously been reported, Ceriporia (Poly- Frisvad et al., 2014). Since different fungi use unique suites porales), Entoloma (Agaricales), and Cheilymenia (Pezizales), as of compounds for chemical defense and nutrient storage well as a sclerotium-forming fungus that could not be iden- (Antibus, 1989; Calvo et al., 2002; Li and Rollins, 2010; Zheng tified to genus (collection MCA3930, Table 1). These structures et al., 2010), sclerotium-forming fungi are excellent targets were also found in a wide range of habitats from cool tem- for the discovery of antibacterial, antifungal, and anti- perate forests in Maine (USA) to lowland tropical forests in herbivore compounds. Secondly, many non-parasitic fungi Guyana. are known to form sclerotia, so it is likely that this life history Although several review articles have discussed morphol- trait is ecologically important for many fungal species and not ogy, function, and diversity of sclerotia, the phylogenetic just for plant pathogens (Chet and Henis, 1975). relationships among the fungi involved were largely unre- During investigations of hypogeous fungi, we encountered solved when these papers were published (Townsend and morphologically variable sclerotia that were not clearly linked Willetts, 1954; Coley-Smith and Cooke, 1971; Chet and Henis, with any known fungus (Fig 1). The wide variation in the 1975). When the affinities of the sclerotium-forming fungi are geography, microhabitats, and morphologies of these scle- viewed within the context of a molecular phylogeny, it is rotia suggested that the sclerotium-forming fungi were not obvious that sclerotium-forming fungi are widely dispersed closely related and differed in their trophic modes. The across the Dikarya. Although more detailed phylogenetic diversity of sclerotia encountered during random sampling analyses are needed to obtain a clear picture of the evolution also suggested the possibility that many fungi form sclerotia of sclerotium formation, we infer that the ability to make in nature but that these structures usually escape detection. sclerotia has probably evolved 14 different times within the The discovery of these varied sclerotia generated several fungi (Fig 2). Our literature review and analysis of new col- questions. First, what are the identities of the unknown lections also suggests that sclerotium formation is infrequent sclerotium-forming fungi found in nature? Second, how many or difficult to observe in some fungal orders (e.g. Dothidiales, unrelated lineages of fungi produce sclerotia? Third, besides ) but common and easy to observe in others plant pathogens, what are the known ecological roles of the (e.g. , Pezizales, Agaricales, Boletales). sclerotium-forming fungi? The sclerotium-forming fungi also represent an extremely To answer these questions, we consulted the published wide trophic diversity. As expected, many sclerotium-forming literature and studied sclerotia collected in nature. To identify fungi are plant pathogens (25 genera) but many other new sclerotium collections, we sequenced ribosomal DNA sclerotium-forming fungi are ectomycorrhizal (11 genera) or (ITS, LSU, and/or SSU) using published protocols and com- saprotrophic (30 genera). The saprotrophs include specialists pared these sequences with GenBank using BLAST searches on distinct substrata such as wood (), humus (Agro- (Table 1) and preliminary phylogenetic analyses (data not cybe), and dung (Cheilymenia). A few genera are also insect shown) (Altschul et al., 1990; Smith and Pfister, 2009; Tedersoo pathogens (Ophiocordyceps), ericoid mycorrhizal (Phialoce- and Smith, 2013). We also surveyed the literature to identify phala), animal pathogens (some Aspergillus), mycoparasites sclerotium-forming fungi by querying Web of Science (www. (Laetisaria), or lichenicolous (Leucogyrophana)(Table 2). Two webofknowledge.com) and Google Scholar (http://scholar. genera (Trechispora, Fibulorhizoctonia) contain putative sapro- google.com/) with key words “sclerotia” and “sclerotium”. To trophs whose sclerotia are tended by termites in an unusual obtain a phylogenetic overview of sclerotium-forming fungi symbiotic relationship that is analogous to brood (Fig 2), we created a database of sclerotium-producing genera (Matsuura and Yashiro, 2010). Several sclerotium-forming by recording one representative species and published refer- fungi, such as purpureum (plant parasite/ ence for each genus reported to form sclerotia (Table 2). These mycoparasite) and arachnoidea (plant pathogen/ sclerotium-producing genera were then mapped onto a lichenicolous), have complex lifecycles that appear to involve

Please cite this article in press as: Smith, et al., How many fungi make sclerotia?, Fungal Ecology (2014), http://dx.doi.org/ 10.1016/j.funeco.2014.08.010 4 laect hsatcei rs s mt,e l,Hwmn ug aeslrta,Fna clg 21) http://dx.doi.org/ (2014), Ecology Fungal sclerotia?, make fungi many How al., et Smith, as: press in 10.1016/j.funeco.2014.08.010 article this cite Please

Table 1 e Collecting data, molecular data, and phylogenetic affiliations of new sclerotia specimens collected in soil, leaf litter, and wood Genus of Inferred Substrate Order Collection Collector and Morphology Collection Most informative GenBank sclerotium- ecology number collection location BLAST match forming and herbarium date fungus

Cheilymenia Saprobe Soil Pezizales Ascoe MES-313 ME Smith, Brown, rounded Mixed forest near DQ220321 Cheilymenia KJ720887 (SSU), (FLAS-F-58920) 3-Aug-09 to irregular Tunk Lake, outside crucipila KJ720888 (LSU) Cherryfield, Maine, (717/734 e 98 %), USA LSU region Unknown Genus ? Soil Amylocorticiales (?) Basidioe MCA3930 ME Smith, Tan, rounded Dicymbe forest in DQ144610 Amyloathelia KJ720886 (FLAS-F-58921) 15-May-10 the Pakaraima crassiuscula (ITS, LSU) Mnts., Guyana (865/1023 e 85 %), ITS and LSU regions Entoloma Ectomycorrhizal Soil and Agaricales Basidioe MES-347 ME Smith, Orange, round Mixed forest with JF908003 Entoloma KJ720892 (LSU), leaf litter (FLAS-F-58922) 18-Dec-09 Dicymbe, Pakaraima platyphylloides KJ720893 (ITS) Mnts., Guyana (603/644 e 94 %), ITS region Ceriporia Saprobe Decayed Polyporales Basidioe MES-332 ME Smith, Tan to cream, Mixed forest, JX644048 Ceriporia KJ720890 (SSU), wood (FLAS-F-58923) 24-Oct-09 irregular Pisgah National Forest, purpurea KJ720891 (LSU), near Marion, (677/704 e 96 %), KJ720889 (ITS) North Carolina, USA LSU region Boletus Ectomycorrhizal Soil and Boletales Basidioe MES-260 (FH) ME Smith, Orange, lobed Angiosperm-dominated EU569236 Boletus sp. FJ480429 (ITS) leaf litter 19-Aug-08 woods, Arlington Great (779/808 e 96 %), Meadows, Arlington, ITS region Massachusetts, USA ..Smith M.E. tal. et How many fungi make sclerotia? 5

Fig 2 e Simplified schematic phylogeny highlighting fungal lineages with sclerotium-forming fungi. Only members of the Dikarya (Ascomycota and Basidiomycota) are shown because no sclerotium-forming fungi have been documented among the early-diverging fungal lineages. Numbers adjacent to fungal orders indicate the number of genera containing at least one sclerotium-forming species. Black circles indicate fungal orders for which all reports of sclerotium formation were obtained from published sources whereas white circles indicate fungal orders for which a new record for sclerotium formation is reported for at least one genus. The schematic phylogeny is based on Hibbett et al. (2007) with phylogenetic positions of new or revised orders inferred from LoBuglio and Pfister (2010), Schoch et al. (2009a, 2009b), Binder et al. (2010), Zhang et al. (2011), Toome et al. (2013), Boehm et al. (2009), Campbell et al. (2009), and Padamsee et al. (2012). The unresolved phylogenetic positions of two sclerotium-forming , Magnaporthe salvinii and Verticilium dahliae, are depicted with broken lines. To reduce the complexity of the figure, some known orders are not shown; asterisks highlight areas of the tree where fungal orders not currently know to form sclerotia have been omitted.

multiple, distantly related host organisms. Still other genera, this trait across the fungal phylogeny along with the diverse such as the putative root endophyte Mattirolomyces and the trophic modes of sclerotium-forming fungi suggests that the putative aphid symbiont , have uncertain trophic biology and evolution of sclerotium formation warrants modes (Brundrett and Kendrick, 1987; Kovacs et al., 2007). additional study. We expect that future research on scle- Taken together, our observations suggest that sclerotium rotium formation will find this feature to be even more widely formation is a more common life history trait among fungi dispersed across the fungal phylogeny than detected here. We than previously recognized. The widespread occurrence of suggest that sclerotium formation is analogous to highly

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Table 2 e Phylogenetic affiliations, trophic modes, and reference information for 85 genera of sclerotium-forming fungi, including three genera that are reported to form sclerotia for the first time: dung-specialized saprobe Cheilymenia (Pezizales), wood decaying Ceriporia (Polyporales), and putatively ectomycorrhizal Entoloma (Agaricales) (this genus contains both saprotrophic and ectomycorrhizal species e Tedersoo and Smith, 2013). One sclerotium collection (MCA3930) found in a tropical rainforest in Guyana putatively belongs to the order Amylocorticiales, but could not be identified to genus based on DNA sequences and has an uncertain trophic mode Genus Species Phylum Lineage Trophic role References

Macrophomina phaseolina A Botryosphaeriales Plant pathogen Papavizas, 1977 Mycosphaerella ligulicola A Capnodiales Plant pathogen Blakeman and Hornby, 1966 Capnobotryella renispora A Capnodiales Plant pathogen Hambleton et al., 2003 Scleroconidioma sphagnicola A Dothideales Plant pathogen Hambleton et al., 2003 Aspergillus flavus A Eurotiales Saprobe, animal Hedayati et al., 2007 pathogen Penecillium sclerotigenum A Eurotiales Saprobe Joshi et al., 1999 Scleromitrula shiraianum A Helotiales Plant pathogen Schumacher and Holst-Jensen, 1997 fuckelinia A Helotiales Plant pathogen Hsiang and Chastagner, 1992 erythronii A Helotiales Plant pathogen Batra and Korf, 1959 carunculoides A Helotiales Plant pathogen Whetzel and Wolf, 1945 tuberosa A Helotiales Plant pathogen Uzuhashi et al., 2010 pyramidalis1 A Helotiales Plant pathogen Grand and Menge, 1974 Kohninia linnaeicola A Helotiales Plant pathogen Holst-Jensen et al., 2004 Martininia panamaensis A Helotiales Saprobe Whetzel, 1942 denisii A Helotiales Plant pathogen Schumacher and Kohn, 1985 azaleae A Helotiales Plant pathogen Weiss, 1940 Redheadia quercus A Helotiales Plant pathogen Suto and Suyama, 2005 Sclerocrana atra A Helotiales Saprobe Samuels and Kohn, 1986 A Helotiales Plant pathogen Kohn, 1979 Septotinia podophyllina A Helotiales Plant pathogen Whetzel, 1945 Streptotinia arisaemae A Helotiales Plant pathogen Whetzel, 1945 gladioli A Helotiales Plant pathogen Whetzel, 1945 Acephala macrosclerotiorum A Helotiales Ectomycorrhizal Munzenberger€ et al., 2009 Phialocephala fortinii A Helotiales Ericoid mycorrhizal Currah et al., 1993 Claviceps purpurea A Hypocreales Plant pathogen Douhan et al., 2008 Ophiocordyceps sinensis A Hypocreales Insect parasite Xing and Guo, 2008 Cylindrocladium crotalariae A Hypocreales Plant pathogen Roth et al., 1979 Cenococcum geophilum A Hysteriales Ectomycorrhizal Douhan and Rizzo, 2005 Verticillium dahliae A Hypocreomycetidae Plant pathogen Tjamos and Fravel, 1995 incertae sedis Magnaporthe salvinii2 A Plant pathogen Cintas and Webster, 2001 incertae sedis crassipes A Pezizales Saprobe Volk and Leonard, 1989 Mattirolomyces terfezioides A Pezizales Root endophyte? Kovacs et al., 2007 Cheilymenia sp. A Pezizales Saprobe This Study Pseudombrophila dentata3 A Pezizales Saprobe Pfister, 1984 Pyronema domesticum A Pezizales Saprobe Moore, 1962 Phymatotrichopsis omnivora A Pezizales Plant pathogen Marek et al., 2009 Wynnea americana A Pezizales Saprobe Pfister, 1979 Coniothyrium glycines4 A Pleosporales Plant pathogen Hartman and Sinclair, 1992 Alternaria brassicae A Pleosporales Plant pathogen Tsuneda and Skoropad, 1977 Paraleptosphaeria orobanches A Pleosporales Plant pathogen de Gruyter et al., 2013 Leptosphaeria sclerotioides5 A Pleosporales Plant pathogen Gray et al., 2008 Colletotrichum coccodes A Sordariales Plant pathogen Blakeman and Hornby, 1966 Sordaria sclerogenia A Sordariales Saprobe Fields and Grear, 1966 Rosellinia necatrix A Xylariales Plant pathogen Gutierrez-Barranquero et al., 2012 Gloeocercospora sorghi6 A Xylariales Plant pathogen Dean, 1968 Leucocoprinus luteus B Agaricales Saprobe Warcup and Talbot, 1962 Pleurotus tuber-regium B Agariacles Saprobe Fasidi and Ekuere, 1993 Coprinus lagopus B Agaricales Saprobe Waters et al., 1975 Cortinarius calochrous B Agaricales Ectomycorrhizal Kernaghan, 2001 Entoloma sp. B Agaricales Ectomycorrhizal? This Study Coprinopsis sclerotiorum B Agaricales Saprobe Keirle et al., 2004 Agrocybe arvalis B Agaricales Saprobe Redhead and Kroeger, 1987 Hebeloma sacchariolens B Agaricales Ectomycorrhizal Ingleby et al., 1990 Hypholoma tuberosum B Agaricales Saprobe Redhead and Kroeger, 1987

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Table 2 e (continued) Genus Species Phylum Lineage Trophic role References

Psilocybe caerulescens B Agaricales Saprobe Redhead and Kroeger, 1987 Stropharia tuberosa B Agaricales Saprobe Redhead and Kroeger, 1987 Collybia tuberosa B Agaricales Saprobe Murrill, 1915 Omphalia lapidescens B Agaricales Saprobe Saito et al., 1992 Rimbachia sp.7 B Agaricales Saprobe (?) Warcup and Talbot, 1962 incarnata B Agaricales Pathogen Matsumoto and Tajimi, 1985 Unknown Genus sp. B Amylocorticiales ? This Study Sclerotium rolfsii8 B Amylocorticiales Saprobe, Binder et al., 2010 plant pathogen Athelia arachnoidea B Atheliales Lichenicolous, Diederich and Lawrey, 2007 plant pathogen Fibularhizoctonia sp. B Atheliales Saprobe, Matsuura, 2006 insect parasite Boletus rubropunctus B Boletales Ectomycorrhizal Smith and Pfister, 2009 Leccinum holopus B Boletales Ectomycorrhizal Muller and Agerer, 1990 aurantiaca B Boletales Saprobe Antibus, 1989 Leucogyrophana lichenicola B Boletales Lichenicolous Diederich and Lawrey, 2007 Boletinellus meruloides B Boletales Insect symbiont? Cotter and Miller, 1985 lividus B Boletales Ectomycorrhizal Agerer et al, 1993 involutus B Boletales Ectomycorrhizal Fox, 1986 sudanicus B Boletales Saprobe Thoen and Ducousso, 1990 Pisolithus tinctorious B Boletales Ectomycorrhizal Piche and Fortin, 1982 Scleroderma verrucosum B Boletales Ectomycorrhizal Ba and Thoen, 1990 Austropaxillus sp. B Boletales Ectomycorrhizal Palfner, 2001 Ceratorhiza hydrophila9 B Cantharelalles Plant pathogen Xu et al., 2010 Rhizoctonia solani10 B Cantharelalles Plant pathogen Cubeta and Vilgalys, 1997 botryohypochnoideum B Corticiales Saprobe Warcup and Talbot, 1962 Laetisaria arvalis B Corticiales Mycoparasite Burdsall et al., 1980 Marchandiomyces lignicola B Corticiales Lichenicolous Larsson, 2007 B Helicobasidiales Plant pathogen, Lutz et al., 2004 mycoparasite Ceriporia sp. B Polyporales Saprobe This Study Lignosus rhinocerus B Polyporales Saprobe Cui et al., 2011 Polyporus mylittae B Polyporales Saprobe Macfarlane et al., 1978 Wolfiporia cocos11 B Polyporales Saprobe Weber, 1929 Trechispora sp. B Trechisporales Saprobe, Matsuura and Yashiro, 2010 insect parasite

Synonyms ¼ 1Cristulariella pyramidalis, 2Sclerotium oryzae, 3Firmaria dentata, 4Dactuliochaeta glycines, 5Phoma scierotioides, 8Athelia rolfsii, 11Poria cocos. Sexual stage ¼ 6Monographella, 9Ceratobasidium, 10Thanatephorus. 7Reported as Leptoglossum sp. 8Binder et al. (2010) showed A. rolfsii is not closely related to Athelia sensu stricto.

adaptive yet massively convergent traits in animals (e.g. Foundation grants DEB-0918591 (TW Henkel) and DEB-3331108 warning coloration, production of shells, flight/gliding) and (R Vilgalys) with permits granted by the Guyana Environ- plants (e.g. thorns, succulents, C4 photosynthesis) but that the mental Protection Agency. MC Aime is acknowledged for her hidden nature of the fungi has concealed the importance of help in photographing and processing sclerotia collection this trait. Lastly, we suspect that the sclerotium-forming fungi MCA3930. contain a veritable treasure trove of interesting secondary metabolites and we suggest that the sclerotium-forming fungi references should be prioritized for genome sequencing and closer metabolomic and ecological study.

Agerer, R., Waller, K., Treu, R., 1993. Die ektomykorrhizen und Acknowledgments sklerotien von . Beiheft zur Zeitschrift fur€ Mykologie 59, 131e140. Funding for ME Smith was provided in part by University of Altschul, S.F., Gish, W., Miller, W., Myers, E.W., Lipman, D.J., 1990. Basic local alignment search tool. Journal of Molecular Biology Florida’s Institute of Food and Agricultural Sciences (IFAS). 215 (3), 403e410. Collecting of sclerotia in New England was made possible via a Antibus, R.K., 1989. Formation and structure of the sclerotia and fellowship provided by the Harvard University Herbaria to ME sclerotium-specific proteins in Hygrophoropsis aurantiaca. Smith. Collecting in Guyana was funded by National Science Mycologia 81 (6), 905e913.

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