Systematics and Evolution, Part A The Mycota

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Boletopsis,Hydnellum,Phellodon,Polyozellus,
Pseudotomentella,Sarcodon, and Thelephora.
This compound also occurs in Boletales and
other orders but nonetheless seems to be a
distinguishing feature of the group (Bresinsky
and Rennschmid 1971 ).
Ecological diversity: most members of The-
lephorales are ECM and are often dominant
components of mycorrhizal communities
(Bruns et al. 1998 ; Tedersoo et al. 2010 ). How-
ever,Lenzitopsisproduces fruiting bodies on
wood of junipers and is reported to produce a
white rot (Zhou and Ko ̃ljalg 2013 ).Amaurodon
is also reported to grow on wood of living trees
(U. Ko ̃ljalg, unpublished) and is presumably
nonmycorrhizal.Amaurodonhas been placed
as the sister group to the remaining Thelephor-
ales, butLenzitopsisis nested within the group,
closely related toTomentellopsis, suggesting
that there have been multiple transitions
between nutritional modes (Larsson 2007b;
Zhou and Ko ̃ljalg 2013 ).
Systematics: the current classification of
Thelephorales (Kirk et al. 2008 ) includes two
families, Thelephoraceae and Bankeraceae.
Donk ( 1964 , p. 247) thought that the similarity
of the Bankeraceae to certain Thelephoraceae
was “an example of extreme convergence,” but
other authors suggested that the two families
were closely related (Ju ̈lich 1981 ; Stalpers 1993 ),
and this has been repeatedly supported by
molecular data (Binder et al. 2005 ; Bruns et al.
1998 ; Larsson 2004 ,2007b; Zhou and Ko ̃ljalg
2013 ). The analysis of Zhou and Ko ̃ljalg ( 2013 )
resolved two nonsister clades corresponding
to Bankeraceae (one including Bankeraand
Phellodonand another containingHydnellum,
Sarcodon, andBoletopsis) and a paraphyletic
assemblage of taxa corresponding to Thele-
phoraceae. However, internal support for
many deep nodes was weak. An in-depth multi-
gene phylogenetic analysis is needed to assess
the classification of the order.


I. Corticiales


Overview: Corticiales K.H. Larsson is a small
order established to accommodate basidiomy-
cetes recognized in recent molecular phyloge-


netic studies and included previously in the
Vuilleminiales (Boidin et al. 1998 ), theDendro-
corticiumclade (Binder and Hibbett 2002 ), and
the corticioid clade (Binder et al. 2005 ; Larsson
et al. 2004). It has been represented by a single
family, Corticiaeae Herter, made up of ca. 29
genera and 136 species (Kirk et al. 2008 ), but
the molecular phylogenetic study of Ghobad-
Nejhad et al. ( 2010 ) recognized three families
and several new genera (discussed subse-
quently). The order is made up of mostly resu-
pinate species that produce smooth
hymenophores, a monomitic hyphal system
with or without clamps, and smooth basidios-
pores, often with pink walls. Many species pro-
duce pink or red basidiomata with a
catahymenium in which young basidia do not
form a palisade but are formed deep within a
layer of hyphidia and then elongate to reach the
hymenial surface, and some have dendrohyphi-
dia; however, there is no morphological synap-
omorphy that characterizes the entire order.
Some species are known only from asexual
stages. One genus of these is the anamorph-
typified Marchandiomyces, which was com-
monly included as a core genus in recent
molecular studies. The genus was originally
established for asexual lichen parasites (Dieder-
ich 1990 ; Etayo and Diederich 1996 ), but sexual
forms are also now known for the group. These
includeMarchandiobasidium aurantiacumas
the teleomorph ofMarchandiomyces aurantia-
cus(Diederich et al. 2003 ) andMarchandiopsis
quercina, a species previously assigned toLae-
ticorticiumorVuilleminiathat was found to be
nested among asexualMarchandiomycesspe-
cies by Ghobad-Nejhad et al. ( 2010 ). The clade
containing Marchandiomyces also includes
described plant pathogens in the teleomorph-
typified genera Laetisariaand Limonomyces,
indicating that sexual–asexual relationships
among these species will require more study.
The type species of the order, Corticium
roseum, may form a bulbil-like anamorph
known asHyphelia rosea; these bulbils are sim-
ilar to those ofLaetisaria and Marchandio-
myces(Eriksson and Ryvarden 1976 ).
Ecological diversity: fungi in Corticiales
exhibit a remarkable range of ecologies,
including saprotrophs, plant pathogens, lichen

400 D.S. Hibbett et al.

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