taxa) are rarely documented from mammal
feces. It is possible that spore dispersal of Phal-
lales is entirely dependent on arthropods.
Systematics: Phallales was described by
Fischer ( 1900 ) with two families, Phallaceae
and Clathraceae. A third family, Claustulaceae,
was added to the order (Cunningham 1931 ;
Ju ̈lich 1981 ; Zeller 1949 ), and this concept has
been accepted for a long time. Miller and Miller
( 1988 ) further expanded the ordinal concept by
including Protophallaceae in the order, but they
also included Hysterangiaceae (now in Hyster-
angiales). Currently the order contains six
families (Phallaceae, Clathraceae, Lysuraceae,
Protophallaceae, Claustulaceae, and Trappea-
ceae), and the monophyly of the order and
each family is strongly supported by multigene
phylogenetic analyses (Hosaka et al. 2006 ).
The basal grades of the order are composed
of Protophallaceae, Claustulaceae, and Trap-
peaceae, all of which exhibit an exclusively
sequestrate habit, indicating that stinkhornlike
fruit bodies are derived morphologies in Phal-
lales (Hosaka et al. 2006 ). Among phylogenies
of Agaricomycetes, Phallales represents the sole
example of an unambiguous transition from
sequestrate to nonsequestrate forms.
A total of 29 genera and ca. 100 species are
currently recorded in the order (Kirk et al.
2008 ), but some genera, such asProtuberaand
Trappea, are polyphyletic, with species in both
Phallales and Hysterangiales (Hosaka et al.
2006 ), and require further taxonomic revision.
Some new genera and species have been
described recently (Cabral et al. 2012 ; Desjardin
and Perry 2009 ). Because the center of diversity
of this order probably lies in the tropics (Miller
and Miller 1988 ) and many such areas have not
been extensively investigated, the number of
taxa in this group will be significantly higher
in the future.
- Gomphales
Overview: the fungi in Gomphales (Ju ̈lich 1981 ;
from the Greekpluglike) have long been recog-
nized as a distinct, highly variable clade of
Agaricomycetes (Bruns et al. 1998 ; Hibbett
and Thorn 2001 ; Hosaka et al. 2006 ; Pine
et al. 1999 ). They are characterized by a wide
range of fruiting body morphologies, from
stalked ramarioid/clavarioid (e.g., Ramaria,
Phaeoclavulina, and Lentariaceae) to club
(Clavariadelphaceae), gilled (Gloeocantharel-
lus), cantharelloid-gomphoid (Gomphus,
Phaeoclavulina, andTurbinellus) (Fig.14.4d),
tooth (Beenakia), resupinate-odontoid (Kavi-
nia) (Fig. 14.4e), all the way to sequestrate
fungi (Gauteriaceae) (Giachini et al. 2010 ; Hos-
aka et al. 2006 ; Humpert et al. 2001 ).
Ecological diversity: members of Gom-
phales show heterogeneity in their ecological
characters. Most species in Beenakiaceae, Len-
tariaceae, Kaviniaceae,Gloeocantharellus, and
Phaeoclavulinaand some species ofRamaria
(e.g.,R. moelleriana,R. stricta, andR. circi-
nans) grow and fruit on woody debris, a trait
that has led to their general categorization as
saprotrophs. The other taxa of the order are
generally considered ECM, and while the nutri-
tional status of many species of Gomphales is
still unknown, the formation of ectomycorrhi-
zae by Turbinellus, Gomphus, and some
Ramariaspecies has been confirmed (Agerer
1996a,b,c,d; Agerer and Iosifidou 2004 ; Agerer
et al. 1998 ; Castellano 1988 ; Griffiths et al. 1991 ;
Masui 1926 , 1927 ; Miller and Miller 1988 ;
Nouhra et al. 2005 ; Rinaldi et al. 2008 ).
Systematics: the taxonomy and systematics
of the Gomphales has traditionally relied on
morphological characters now known to be
subject to parallel evolution and phenotypic
plasticity (Moncalvo et al. 2000 ). As a family,
Gomphaceae has traditionally been classified
within the Aphyllophorales, along with dis-
tantly related taxa such as Cantharellaceae,
Ganodermataceae, and Polyporaceae (Donk
1964 ). The phylogenetic relationships of mem-
bers of the order Gomphales, including its
monophyly, have been estimated using mole-
cular data (Giachini et al. 2010 ; Hosaka et al.
2006 ; Humpert et al. 2001 ), which revealed that
gomphoid fungi are closely related to
taxa in Geastrales, Hysterangiales, and Phallales
in the subclass Phallomycetidae (Colgan et al.
1997 ; Giachini et al. 2010 ; Hibbett et al.1997a;
Hosaka et al. 2006 ; Humpert et al. 2001 ;
Pine et al. 1999 ). Currently, the order encom-
passes six well-supported families, namelyAgaricomycetes 393