pathogens, and lichenized species (Lawrey et al.
2008 ). The basal position of saprotrophic spe-
cies in Vuilleminiaceae and Punctulariaceae
would indicate that this is the ancestral condi-
tion for the order, but the most derived clade,
Corticiaceae, contains a complex mixture of
ecological forms, which suggests an unusual
tendency for ecological transitions (Ghobad-
Nejhad et al. 2010 ; Lawrey et al. 2008 ). Ecology
can sometimes be used to characterize genera,
but most clades in this order have mixtures of
nutritional modes. Entirely saprotrophic
genera includeCorticium,Giulia, andGalzinia,
butErythricium, which is mostly saprotrophic,
also includes the plant pathogenErythricium
salmonicolor.
Most species ofLimonomyces,Laetisaria,
andWaiteaare plant pathogens or endophytes
but are able to persist in the field as saprotrophs
(Andjic et al. 2005 ; Burdsall 1979 ; Burdsall et al.
1980 ; Stalpers and Loerakker 1982 ). However,
Laetisariaalso includes the lichen parasiteL.
lichenicola(Diederich et al. 2011 ).Marchandio-
myces, originally described for lichen parasites
(Diederich 1990 ; Etayo and Diederich 1996 ), is
now known to include saprotrophic and folii-
colous species (Diederich and Lawrey 2007 ;
Lawrey et al. 2007 , 2008 ). There is also one
lichen-forming species, Marchandiomphalina
foliacea(Lawrey et al. 2008 ; Palice et al. 2005 ),
which seems to be most closely related to the
lichen parasiteMarchandiobasidium aurantia-
cumand the saprotrophicErythricium laetum.
Lichen mutualisms have evolved independently
in Basidiomycota at least five times (Diederich
and Lawrey 2007 ; Diederich et al. 2003 , 2011 ;
Ertz et al. 2008 ; Fischer et al. 2007 ; Hodkinson
et al. 2012 ; Lawrey et al. 2007 , 2008 , 2009 ;
Nelsen et al. 2007 ; Redhead et al. 2002 ).
Marchandiomphalina foliacea is the only
described lichen species in Corticiales. It
forms a foliose thallus structure and asexual
goniocysts resembling soredia, but no sexual
stages are known (Jørgensen 1989 ).
Systematics: the monophyly of Corticiales
is strongly supported by molecular phyloge-
nies, mostly based on rRNA gene sequences
(Binder et al. 2005 ; Boidin et al. 1998 ; DePriest
et al. 2005 ; Diederich et al. 2011 ; Ghobad-
Nejhad et al. 2010 ; Hibbett et al. 2007 ; Langer
2002 ; Larsson 2007b; Larsson et al. 2004;
Lawrey et al. 2008 ). The sister group of
Corticiales seems, on the basis of many of
these studies, to be the order Gloeophyllales.
One species of Corticiales, Punctularia
strigoso-zonata, has been subject to whole-
genome sequencing; phylogenomic analyses
suggest that it is in a clade that also includes
Gloeophyllales and Jaapiales (Fig. 14.1). A
recent attempt to produce an infraordinal clas-
sification of Corticiales is based on a molecular
phylogeny using nuc-lsu rRNA sequences
(Ghobad-Nejhad et al. 2010 ). This analysis
resolved three groups that were recognized at
the family level using the existing names,
Vuilleminiaceae, Punctulariaceae, and Corticia-
ceae. Until more sequences become available
and more specimens sequenced, this represents
the best current hypothesis for a family
classification of the order.VuilleminiaceaeMaire ex Lotsy: this clade contains
saprotrophic species that develop dendrohyphidia and
produce clamps and generally allantoid spores and
gelatinous fruiting bodies. It includes the generaVuil-
leminiaandCytidiaand a new genus,Australovuille-
minia (for Vuilleminia coccinea). Based on their
molecular phylogeny and incompatibility crossing
tests, Ghobad-Nejhad et al. ( 2010 ) found that the so-
called coreVuilleminiaspecies (V. macrospora,V.
pseudocystidia,V. alni,V. comedens,V. megalospora)
form a monophyletic group. These species are all dec-
orticating, produce a gelatinous fruiting body, and
exhibit a unique 13 bp insertion in the ITS2. Other
describedVuilleminiaspecies (V. cystidiataandV.
macrospora) were recovered in theVuilleminiaclade
but outside of the coreVuilleminiaclade. Other species
were recovered outside of theVuilleminiaclade and
reassigned to new genera, includingV.(Punctulariop-
sis)obduscensandV.(Punctulariopsis)subglobispora,
which were recovered in thePunctulariaclade, and
Vuilleminia (Marchandiopsis) quercina, which was
recovered in theCorticiumclade.
PunctulariaceaeDonk: the family introduced by
Donk ( 1964 ) was intended to separatePunctulariaspe-
cies from other corticioid fungi in the Aphyllophorales,
but few classifications recognized it, placing most cor-
ticioid species in the Corticiaeae. In the nuc-lsu rRNA
phylogeny of Ghobad-Nejhad et al. ( 2010 ), this is a
clade of saprotrophic species that produces clamps
and ellipsoid spores and includes the generaPunctu-
lariaandDendrocorticiumand a new genus,Punctular-
iopsis(forVuilleminia subglobisporaandVuilleminia
obducens). These species cause a vigorous white rot
compared to species inCorticium.Agaricomycetes 401