How many fungi make sclerotia?

Abstract

Most fungi produce some type of durable microscopic structure such as a spore that is important for dispersal and/or survival under adverse conditions, but many species also produce dense aggregations of tissue called sclerotia. These structures help fungi to survive challenging conditions such as freezing, desiccation, microbial attack, or the absence of a host. During studies of hypogeous fungi we encountered morphologically distinct sclerotia in nature that were not linked with a known fungus. These observations suggested that many unrelated fungi with diverse trophic modes may form sclerotia, but that these structures have been overlooked. To identify the phylogenetic affiliations and trophic modes of sclerotium-forming fungi, we conducted a literature review and sequenced DNA from fresh sclerotium collections. We found that sclerotium-forming fungi are ecologically diverse and phylogenetically dispersed among 85 genera in 20 orders of Dikarya, suggesting that the ability to form sclerotia probably evolved ≥14 different times in fungi.

Full text

Short Communication
How many fungi make sclerotia?
Matthew E. SMITH
a,
*, Terry W. HENKEL
b
, Jeffrey A. ROLLINS
a
a
University of Florida, Department of Plant Pathology, Gainesville, FL 32611-0680, USA
b
Humboldt State University of Florida, Department of Biological Sciences, Arcata, CA 95521, USA
article info
Article history:
Received 25 April 2014
Revision received 23 July 2014
Accepted 28 July 2014
Available online -
Corresponding editor:
Dr. Jean Lodge
Keywords:
Chemical defense
Ectomycorrhizal
Plant pathogens
Saprotrophic
Sclerotium
abstract
Most fungi produce some type of durable microscopic structure such as a spore that is
important for dispersal and/or survival under adverse conditions, but many species also
produce dense aggregations of tissue called sclerotia. These structures help fungi to survive
challenging conditions such as freezing, desiccation, microbial attack, or the absence of a
host. During studies of hypogeous fungi we encountered morphologically distinct sclerotia
in nature that were not linked with a known fungus. These observations suggested that
many unrelated fungi with diverse trophic modes may form sclerotia, but that these
structures have been overlooked. To identify the phylogenetic affiliations and trophic
modes of sclerotium-forming fungi, we conducted a literature review and sequenced DNA
from fresh sclerotium collections. We found that sclerotium-forming fungi are ecologically
diverse and phylogenetically dispersed among 85 genera in 20 orders of Dikarya, suggesting
that the ability to form sclerotia probably evolved 14 different times in fungi.
ª2014 Elsevier Ltd and The British Mycological Society. All rights reserved.
Fungi are among the most diverse lineages of eukaryotes with
an estimated 5.1 million species (Blackwell, 2011). They are the
principle saprotrophs in most terrestrial biomes and play
important ecological and economic roles as plant pathogens
and mutualists. Fungi are found in all terrestrial ecosystems
and they use a variety of strategies to colonize appropriate
substrata and survive unfavorable conditions (Blackwell,
2011). They have significant impacts on the biology of plants
because they are the most economically significant plant
pathogens, serve as mycorrhizal and endophytic symbionts,
and act as key players in nutrient cycles (Schumann, 1991;
Rodriguez et al., 2009; Hobbie and Hogberg, 2012). Two fun-
gal phyla, Basidiomycota and Ascomycota, comprise the
subkingdom Dikarya, a diverse group with ca. 100,000
described species (James et al., 2006). Most Dikarya share key
features such as a hyphal thallus, non-flagellated cells, and
the production of spores (Stajich et al., 2009). However,
because of their cryptic lifestyles within environments such
as plants and soil, the ecology and evolutionary history of
many fungi remains poorly understood.
Almost all fungi produce some type of durable, quiescent
microscopic structure such as a spore that is important for
dispersal and/or survival under adverse conditions (Stajich
et al., 2009). However, some fungi also produce dense aggre-
gations of fungal tissue called sclerotia (Willetts, 1971). These
persistent structures help fungi to survive challenging con-
ditions such as freezing temperatures, desiccation, microbial
attack, or the long-term absence of a host (Townsend and
Willetts, 1954; Coley-Smith and Cooke, 1971). Sclerotia are
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).
available at www.sciencedirect.com
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http://dx.doi.org/10.1016/j.funeco.2014.08.010
1754-5048/ª2014 Elsevier Ltd and The British Mycological Society. All rights reserved.
fungal ecology xxx (2014) 1e10
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

melanized rind enclosing compact, undifferentiated hyphae
while others lack a rind (Willetts, 1971). Some species make
round, determinate sclerotia but others have indeterminate
forms where the shape and size are influenced by resources
and environmental conditions (Chet and Henis, 1975). Some
sclerotia are produced inside of host tissues; Claviceps purpurea
produces sclerotia in grass florets after it has destroyed the
plant cells (Douhan et al., 2008) and Ophiocordyceps sinensis
colonizes caterpillars and transforms their tissues into a
sclerotium (Xing and Guo, 2008). In contrast, some fungi make
sclerotia that are spatially separated from hosts (e.g. Phyma-
totrichopsis omnivora forms sclerotia deep in soil eLyda, 1984).
Sclerotia also range in size from “microsclerotia” <1mm
across, as in the plant pathogen Macrophominia phaseolina
(Short and Wyllie, 1978), to the massive sclerotia of Polyporus
mylittae that reach over 40 cm in diameter (Macfarlane et al.,
1978). Sclerotia putatively serve a resource-storage and sur-
vival role in all sclerotium-forming fungi. However, some
fungi such as Sclerotinia sclerotiorum produce sexual fruiting
structures directly on sclerotia (Bolton et al., 2006) whereas
others such as Pteromyces flavus (¼Aspergillus flavus) produce
fruiting bodies within sclerotia (Horn et al., 2009). Still others,
such as Boletus rubropunctus, produce fruit bodies and sclerotia
at different times or in different places (Smith and Pfister,
2009).
Although sclerotia have been documented in several fun-
gal lineages, sclerotium formation is primarily recognized as a
key life history trait in several necrotrophic plant pathogens
(e.g. Sclerotium rolfsii,Rhizoctonia solani,M. phaseolina,P. omni-
vora,S. sclerotiorum). Collectively, these devastating host gen-
eralist pathogens are responsible for hundreds of millions of
dollars in global crop losses annually (Aycock, 1966; Parmeter,
1971; Purdy, 1979; Mulrean et al., 1984). For example, S. scle-
rotiorum and S. rolfsii each attack >400 plant species, including
major crops such as peanuts, potatoes, and soybeans, and can
cause up to 100 % yield losses (Jenkins and Averre, 1986;
Fig 1 eMorphologically 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; Agaricales) from soil in a tropical rainforest dominated by leguminous
ectomycorrhizal trees, Guyana; (C) Cheilymenia sp. (MES 313; Pezizales) 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; Boletales) 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.
2 M.E. Smith et al.
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

Bowen et al., 1992; Cintas and Webster, 2001). For these and
other sclerotium-forming pathogens, survival is tightly linked
with sclerotium formation so sclerotia eradication is critical
for disease control (Coley-Smith and Cooke, 1971). Fur-
thermore, the ecology of these fungi cannot be fully under-
stood without understanding the biology of sclerotium
formation.
Although management of serious plant pathogens is an
important rationale for studying sclerotium formation, there
are nevertheless several other compelling reasons. First,
many sclerotia lie dormant in soil, leaf litter, or wood for
months, so they must survive attacks from a wide variety of
natural enemies, including bacteria, other fungi, and inver-
tebrates (Willetts, 1971; Papavizas, 1977; Matsumoto and
Tajimi, 1985). The mechanisms that allow sclerotia to sur-
vive in soil despite ongoing biotic assault are not well known,
but evidence from well-studied species (e.g. S. sclerotiorum, C.
purpurea) suggests that most sclerotia contain biologically
active secondary metabolites (Morrall et al., 1978; Demain,
1999; Schardl et al., 2006; Ikewuchi and Ikewuchi, 2008;
Frisvad et al., 2014). Since different fungi use unique suites
of compounds for chemical defense and nutrient storage
(Antibus, 1989; Calvo et al., 2002; Li and Rollins, 2010; Zheng
et al., 2010), sclerotium-forming fungi are excellent targets
for the discovery of antibacterial, antifungal, and anti-
herbivore compounds. Secondly, many non-parasitic fungi
are known to form sclerotia, so it is likely that this life history
trait is ecologically important for many fungal species and not
just for plant pathogens (Chet and Henis, 1975).
During investigations of hypogeous fungi, we encountered
morphologically variable sclerotia that were not clearly linked
with any known fungus (Fig 1). The wide variation in the
geography, microhabitats, and morphologies of these scle-
rotia suggested that the sclerotium-forming fungi were not
closely related and differed in their trophic modes. The
diversity of sclerotia encountered during random sampling
also suggested the possibility that many fungi form sclerotia
in nature but that these structures usually escape detection.
The discovery of these varied sclerotia generated several
questions. First, what are the identities of the unknown
sclerotium-forming fungi found in nature? Second, how many
unrelated lineages of fungi produce sclerotia? Third, besides
plant pathogens, what are the known ecological roles of the
sclerotium-forming fungi?
To answer these questions, we consulted the published
literature and studied sclerotia collected in nature. To identify
new sclerotium collections, we sequenced ribosomal DNA
(ITS, LSU, and/or SSU) using published protocols and com-
pared these sequences with GenBank using BLAST searches
(Table 1) and preliminary phylogenetic analyses (data not
shown) (Altschul et al., 1990; Smith and Pfister, 2009; Tedersoo
and Smith, 2013). We also surveyed the literature to identify
sclerotium-forming fungi by querying Web of Science (www.
webofknowledge.com) and Google Scholar (http://scholar.
google.com/) with key words “sclerotia” and “sclerotium”. To
obtain a phylogenetic overview of sclerotium-forming fungi
(Fig 2), we created a database of sclerotium-producing genera
by recording one representative species and published refer-
ence for each genus reported to form sclerotia (Table 2). These
sclerotium-producing genera were then mapped onto a
schematic phylogeny based on Hibbett et al. (2007) with phy-
logenetic 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). All but two fungal species, Magnaporthe salvinii and
Verticilium dahliae, were easily resolved at the ordinal level
based on data from published references (Table 2) and Index
Fungorum (www.indexfungorum.org/).
We documented reports of sclerotium formation in species
from 85 fungal genera in at least 20 orders of Basidiomycota
and Ascomycota (Table 2,Fig 2). Since only one representative
sclerotium-forming species from each genus was recorded,
we cannot accurately estimate the number of sclerotium-
forming species. However, we observed that many genera
with one sclerotium-forming species also contain others.
Also, despite our limited sampling of sclerotia, we found a
wide diversity of sclerotium-forming fungi in nature and
documented at least three genera for which sclerotium for-
mation had not previously been reported, Ceriporia (Poly-
porales), Entoloma (Agaricales), and Cheilymenia (Pezizales), as
well as a sclerotium-forming fungus that could not be iden-
tified to genus (collection MCA3930, Table 1). These structures
were also found in a wide range of habitats from cool tem-
perate forests in Maine (USA) to lowland tropical forests in
Guyana.
Although several review articles have discussed morphol-
ogy, function, and diversity of sclerotia, the phylogenetic
relationships among the fungi involved were largely unre-
solved when these papers were published (Townsend and
Willetts, 1954; Coley-Smith and Cooke, 1971; Chet and Henis,
1975). When the affinities of the sclerotium-forming fungi are
viewed within the context of a molecular phylogeny, it is
obvious that sclerotium-forming fungi are widely dispersed
across the Dikarya. Although more detailed phylogenetic
analyses are needed to obtain a clear picture of the evolution
of sclerotium formation, we infer that the ability to make
sclerotia has probably evolved 14 different times within the
fungi (Fig 2). Our literature review and analysis of new col-
lections also suggests that sclerotium formation is infrequent
or difficult to observe in some fungal orders (e.g. Dothidiales,
Helicobasidiales) but common and easy to observe in others
(e.g. Helotiales, Pezizales, Agaricales, Boletales).
The sclerotium-forming fungi also represent an extremely
wide trophic diversity. As expected, many sclerotium-forming
fungi are plant pathogens (25 genera) but many other
sclerotium-forming fungi are ectomycorrhizal (11 genera) or
saprotrophic (30 genera). The saprotrophs include specialists
on distinct substrata such as wood (Pleurotus), humus (Agro-
cybe), and dung (Cheilymenia). A few genera are also insect
pathogens (Ophiocordyceps), ericoid mycorrhizal (Phialoce-
phala), animal pathogens (some Aspergillus), mycoparasites
(Laetisaria), or lichenicolous (Leucogyrophana)(Table 2). Two
genera (Trechispora, Fibulorhizoctonia) contain putative sapro-
trophs whose sclerotia are tended by termites in an unusual
symbiotic relationship that is analogous to brood parasitism
(Matsuura and Yashiro, 2010). Several sclerotium-forming
fungi, such as Helicobasidium purpureum (plant parasite/
mycoparasite) and Athelia arachnoidea (plant pathogen/
lichenicolous), have complex lifecycles that appear to involve
How many fungi make sclerotia? 3
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10.1016/j.funeco.2014.08.010

Table 1 eCollecting data, molecular data, and phylogenetic affiliations of new sclerotia specimens collected in soil, leaf litter, and wood
Genus of
sclerotium-
forming
fungus
Inferred
ecology
Substrate Order Phylum Collection
number
and herbarium
Collector and
collection
date
Morphology Collection
location
Most informative
BLAST match
GenBank
Cheilymenia Saprobe Soil Pezizales AscoeMES-313
(FLAS-F-58920)
ME Smith,
3-Aug-09
Brown, rounded
to irregular
Mixed forest near
Tunk Lake, outside
Cherryfield, Maine,
USA
DQ220321 Cheilymenia
crucipila
(717/734 e98 %),
LSU region
KJ720887 (SSU),
KJ720888 (LSU)
Unknown Genus ? Soil Amylocorticiales (?) BasidioeMCA3930
(FLAS-F-58921)
ME Smith,
15-May-10
Tan, rounded Dicymbe forest in
the Pakaraima
Mnts., Guyana
DQ144610 Amyloathelia
crassiuscula
(865/1023 e85 %), ITS
and LSU regions
KJ720886
(ITS, LSU)
Entoloma Ectomycorrhizal Soil and
leaf litter
Agaricales BasidioeMES-347
(FLAS-F-58922)
ME Smith,
18-Dec-09
Orange, round Mixed forest with
Dicymbe, Pakaraima
Mnts., Guyana
JF908003 Entoloma
platyphylloides
(603/644 e94 %),
ITS region
KJ720892 (LSU),
KJ720893 (ITS)
Ceriporia Saprobe Decayed
wood
Polyporales BasidioeMES-332
(FLAS-F-58923)
ME Smith,
24-Oct-09
Tan to cream,
irregular
Mixed forest,
Pisgah National Forest,
near Marion,
North Carolina, USA
JX644048 Ceriporia
purpurea
(677/704 e96 %),
LSU region
KJ720890 (SSU),
KJ720891 (LSU),
KJ720889 (ITS)
Boletus Ectomycorrhizal Soil and
leaf litter
Boletales BasidioeMES-260 (FH) ME Smith,
19-Aug-08
Orange, lobed Angiosperm-dominated
woods, Arlington Great
Meadows, Arlington,
Massachusetts, USA
EU569236 Boletus sp.
(779/808 e96 %),
ITS region
FJ480429 (ITS)
4 M.E. Smith et al.
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multiple, distantly related host organisms. Still other genera,
such as the putative root endophyte Mattirolomyces and the
putative aphid symbiont Boletinellus, have uncertain trophic
modes (Brundrett and Kendrick, 1987; Kovacs et al., 2007).
Taken together, our observations suggest that sclerotium
formation is a more common life history trait among fungi
than previously recognized. The widespread occurrence of
this trait across the fungal phylogeny along with the diverse
trophic modes of sclerotium-forming fungi suggests that the
biology and evolution of sclerotium formation warrants
additional study. We expect that future research on scle-
rotium formation will find this feature to be even more widely
dispersed across the fungal phylogeny than detected here. We
suggest that sclerotium formation is analogous to highly
Fig 2 eSimplified 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 Sordariomycetes, 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.
How many fungi make sclerotia? 5
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Table 2 ePhylogenetic 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 eTedersoo 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
pathogen
Hedayati et al., 2007
Penecillium sclerotigenum A Eurotiales Saprobe Joshi et al., 1999
Scleromitrula shiraianum A Helotiales Plant pathogen Schumacher and Holst-Jensen,
1997
Botryotinia fuckelinia A Helotiales Plant pathogen Hsiang and Chastagner, 1992
Ciborinia erythronii A Helotiales Plant pathogen Batra and Korf, 1959
Ciboria carunculoides A Helotiales Plant pathogen Whetzel and Wolf, 1945
Dumontinia tuberosa A Helotiales Plant pathogen Uzuhashi et al., 2010
Grovesinia pyramidalis
1
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
Myriosclerotinia denisii A Helotiales Plant pathogen Schumacher and Kohn, 1985
Ovulinia 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
Sclerotinia sclerotiorum A Helotiales Plant pathogen Kohn, 1979
Septotinia podophyllina A Helotiales Plant pathogen Whetzel, 1945
Streptotinia arisaemae A Helotiales Plant pathogen Whetzel, 1945
Stromatinia gladioli A Helotiales Plant pathogen Whetzel, 1945
Acephala macrosclerotiorum A Helotiales Ectomycorrhizal M€
unzenberger 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
incertae sedis
Plant pathogen Tjamos and Fravel, 1995
Magnaporthe salvinii
2
A Sordariomycetidae
incertae sedis
Plant pathogen Cintas and Webster, 2001
Morchella crassipes A Pezizales Saprobe Volk and Leonard, 1989
Mattirolomyces terfezioides A Pezizales Root endophyte? Kov 
acs et al., 2007
Cheilymenia sp. A Pezizales Saprobe This Study
Pseudombrophila dentata
3
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 glycines
4
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 sclerotioides
5
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 Guti
errez-Barranquero et al.,
2012
Gloeocercospora sorghi
6
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
6 M.E. Smith et al.
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

adaptive yet massively convergent traits in animals (e.g.
warning coloration, production of shells, flight/gliding) and
plants (e.g. thorns, succulents, C4 photosynthesis) but that the
hidden nature of the fungi has concealed the importance of
this trait. Lastly, we suspect that the sclerotium-forming fungi
contain a veritable treasure trove of interesting secondary
metabolites and we suggest that the sclerotium-forming fungi
should be prioritized for genome sequencing and closer
metabolomic and ecological study.
Acknowledgments
Funding for ME Smith was provided in part by University of
Florida’s Institute of Food and Agricultural Sciences (IFAS).
Collecting of sclerotia in New England was made possible via a
fellowship provided by the Harvard University Herbaria to ME
Smith. Collecting in Guyana was funded by National Science
Foundation grants DEB-0918591 (TW Henkel) and DEB-3331108
(R Vilgalys) with permits granted by the Guyana Environ-
mental Protection Agency. MC Aime is acknowledged for her
help in photographing and processing sclerotia collection
MCA3930.
references
Agerer, R., Waller, K., Treu, R., 1993. Die ektomykorrhizen und
sklerotien von Gyrodon lividus.Beiheft zur Zeitschrift f€
ur
Mykologie 59, 131e140.
Altschul, S.F., Gish, W., Miller, W., Myers, E.W., Lipman, D.J., 1990.
Basic local alignment search tool. Journal of Molecular Biology
215 (3), 403e410.
Antibus, R.K., 1989. Formation and structure of the sclerotia and
sclerotium-specific proteins in Hygrophoropsis aurantiaca.
Mycologia 81 (6), 905e913.
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
Typhula incarnata B Agaricales Pathogen Matsumoto and Tajimi, 1985
Unknown Genus sp. B Amylocorticiales ? This Study
Sclerotium rolfsii
8
B Amylocorticiales Saprobe,
plant pathogen
Binder et al., 2010
Athelia arachnoidea B Atheliales Lichenicolous,
plant pathogen
Diederich and Lawrey, 2007
Fibularhizoctonia sp. B Atheliales Saprobe,
insect parasite
Matsuura, 2006
Boletus rubropunctus B Boletales Ectomycorrhizal Smith and Pfister, 2009
Leccinum holopus B Boletales Ectomycorrhizal Muller and Agerer, 1990
Hygrophoropsis 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
Gyrodon lividus B Boletales Ectomycorrhizal Agerer et al, 1993
Paxillus involutus B Boletales Ectomycorrhizal Fox, 1986
Phlebopus 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 hydrophila
9
B Cantharelalles Plant pathogen Xu et al., 2010
Rhizoctonia solani
10
B Cantharelalles Plant pathogen Cubeta and Vilgalys, 1997
Corticium botryohypochnoideum B Corticiales Saprobe Warcup and Talbot, 1962
Laetisaria arvalis B Corticiales Mycoparasite Burdsall et al., 1980
Marchandiomyces lignicola B Corticiales Lichenicolous Larsson, 2007
Helicobasidium purpureum B Helicobasidiales Plant pathogen,
mycoparasite
Lutz et al., 2004
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 cocos
11
B Polyporales Saprobe Weber, 1929
Trechispora sp. B Trechisporales Saprobe,
insect parasite
Matsuura and Yashiro, 2010
Synonyms ¼
1
Cristulariella pyramidalis,
2
Sclerotium oryzae,
3
Firmaria dentata,
4
Dactuliochaeta glycines,
5
Phoma scierotioides,
8
Athelia rolfsii,
11
Poria
cocos.
Sexual stage ¼
6
Monographella,
9
Ceratobasidium,
10
Thanatephorus.
7
Reported as Leptoglossum sp.
8
Binder et al. (2010) showed A. rolfsii is not closely related to Athelia sensu stricto.
How many fungi make sclerotia? 7
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10.1016/j.funeco.2014.08.010

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