also been called Coniophoraceae-type rot
(Ka ̈mmerer et al. 1985 ; Nilsson and Ginns
1979 ) to separate it from other brown-rot
types. Most brown-rot-causing species contrib-
ute to carbon sequestration in conifer forests
(Tapinella,Pseudomerulius), but a few have
specialized on human-built timber environ-
ments. The so-called cellar fungusConiophora
puteanaand especially the so-called dry rot
fungus Serpula lacrymans cause significant
damage in wooden building structures
(Schmidt and Kebernik 1989 ; Schmidt et al.
2002 ), and the ecological diversification and
structure of geographical lineages of these
aggressive decayers have been studied in detail
(Eastwood et al. 2011 ; Kauserud et al.2007a,b;
Skrede et al. 2011 ; Watkinson and Eastwood
2012 ). Serpulaceae is also a prime example of
transitions from brown-rot to ectomycorrhiza
associated with major morphological changes.
Austropaxillusspecies (pileate-stipitate fruiting
bodies with gilled hymenophores) andGymno-
paxillusspecies (gasteroid) are derived from
withinSerpulaand form ectomycorrhizae with
Eucalyptusand Nothofagus(Bresinsky et al.
1999 ; Jarosch 2001 ; Skrede et al. 2011 ).
Approximately 90 % of species in Boletales
are involved in ECM symbioses, particularly
with Betulaceae, Caesalpiniaceae, Dipterocarpa-
ceae, Fagaceae, Myrtaceae, Nothofagaceae, Pina-
ceae, and Salicaceae, or in arbutoid mycorrhiza
with Ericaceae (Newman and Reddell 1987 ;
Rinaldi et al. 2008 ; Tedersoo et al. 2010 ).Boletus
andLeccinumspp. show an increased tendency
to associate with specific hosts; for example,
Leccinum scabrumforms ectomycorrhizae with
Betula (Singer 1967 ). Suillus, Gomphidius,
Chroogomphus,andRhizopogon spp. in the
suborder Suillineae are almost exclusively asso-
ciated with Pinaceae, which is probably the old-
est clade of ECM partners for Boletales (Hibbett
and Matheny 2009 ). Most ECM species are
placed among the Boletaceae (roughly 400 plus
species), which include highly prized edibles
such asBoletus edulis(porcini).
Mycoparasitesin Boletales represent tran-
sitions from the ECM lifestyle (Binder and
Hibbett 2006 ) and have evolved at least twice
independently.GomphidiusandChroogomphus
spp. are capable of parasitizing the established
ectomycorrhizae of the closely relatedSuillus
and Rhizopogon by penetrating their rhizo-
morphs (Agerer 1990 , 1999 ; Miller 1964 ; Olsson
et al. 2000 ), thereby circumventing competition
for host plants (Binder and Hibbett 2006 ).Pseu-
doboletus parasiticusin Boletaceae produces its
fruiting bodies onScleroderma citrinum(a gas-
teroid bolete) while eroding the gleba of the
host (Binder and Hibbett 2006 ). Other putative
mycoparasites from the basal lineages of Bole-
taceae include the sister taxaBuchwaldoboletus
andChalciporus(Nuhn et al. 2013 ).
Systematics: Boletales includes five subor-
ders that have been described based on dispa-
rate characteristics and methods: Boletineae,
Suillineae, Sclerodermatineae, Tapinellineae,
and Coniophorineae (Binder and Hibbett
2006 ). Boletineae was first introduced by
Gilbert ( 1931 ) using fruiting body morphology
and spore shape as distinctive characteristics.
This suborder included all species with tubular
hymenophores at that time and a few species
with gilled hymenophores. Suillineae was sepa-
rated later from Boletineae based on unique
pigments occurring in this group (Besl and
Bresinsky 1997 ). In addition, numerous resupi-
nate and paxilloid taxa producing a brown rot
were known to be closely related to Boletales
based on their pigments (Besl et al. 1986 ), but
they remainedincertae sedis. The morphology
of belowground rhizomorphs helped to for-
mally place these taxa in Tapinellineae and
Coniophorineae (Agerer 1999 ), which was sup-
ported by early major studies using DNA
sequences (Bruns et al. 1998 ; Kretzer and
Bruns 1999 ). Sclerodermatineae was described
based on nuc-lsu rRNA sequences (Binder and
Bresinsky 2002 ) integrating the gasteroid Scler-
odermatales into Boletales.
Resupinate taxaare still a source of taxo-
nomic uncertainty in Tapinellineae and Con-
iophorineae, particularly concerning the
polyphyletic genus Leucogyrophana (Binder
et al. 2010 ; Jarosch and Besl 2001 ). Conio-
phorineae, including three clades ofLeucogyr-
ophana, has been resolved as a monophyletic
group, but without significant statistical sup-
port (Binder et al. 2010 ). Together, Boletineae,
Sclerodermatineae, and Suillineae form the
largest clade, including the overwhelming408 D.S. Hibbett et al.