Beenakiaceae (Beenakia,KaviniaandRamari-
cium), Clavariadelphaceae (Clavariadelphus),
Gautieriaceae (Gautieria), Gomphaceae (Gloeo-
cantharellus, Gomphus, Phaeoclavulina, and
Turbinellus), Lentariaceae (Lentaria), and
Ramariaceae (Ramaria), distributed within 18
genera and ca. 336 species (Kirk et al. 2008 ).
Despite their macromorphological varia-
tion, the members of the order share a number
of microscopic and macrochemical characters,
including cyanophilic spore ornamentation,
chiastic basidia, similar hyphal construction,
and positive hymenial reaction to ferric sulfate
(Donk 1961 , 1964 ; Eriksson 1954 ; Humpert
et al. 2001 ; Petersen1971b; Villegas et al. 1999 ,
2005 ). In the studies of Hosaka et al. ( 2006 ),
both Bayesian and parsimony analyses showed
strong support for the monophyly of the Phal-
lomycetidae. Even though no definitive synapo-
morphies have been identified for this
gomphoid-phalloid clade, some potential syna-
pomorphic characters, including rhizomorph
morphology (presence of ampullate hyphae
and acanthohypha), pistillarin content, and
structures of septal pore cap, have been pro-
posed (Agerer and Iosifidou 2004 ; Hibbett and
Thorn 2001 ). In addition, some members of the
gomphoid-phalloid clade, such asGautieria,
Hysterangium, Ramaria, and Geastrum, are
known to produce thick hyphal mats in soil
(Agerer and Iosifidou 2004 ; Nouhra et al.
2005 ; Sunhede 1989 ). Although most of these
characters are not exclusive to the gomphoid-
phalloid fungi, the yellowish filled acanthocys-
tidia and associated “exuded drops of pig-
ments” have been reported only from the
gomphoid-phalloid fungi [e.g.,Geastrum,Gom-
phus,Phallogaster, andRamaria(Agerer and
Iosifidou 2004 )].
- Hysterangiales
Overview: this group has long been considered
a sequestrate (trufflelike) relative of stinkhorns
(Phallales). The ordinal status was accepted by
some authors (Hosaka et al. 2006 ;Ju ̈lich 1981 ;
Zeller 1939 ), but others have included the
group in the order Phallales (Kirk et al. 2008 ;
Miller and Miller 1988 ). The order contains
exclusively sequestrate taxa with hypogeous
fruit bodies (e.g.,Hysterangium,Mesophellia,
Austrogautieria) (Fig.14.4c), but taxa of epige-
ous habit (e.g.,Gallacea,Phallogaster) are also
known, and they often expose a gleba at matu-
rity (Castellano and Beever 1994 ). Most taxa are
characterized as having gelatinous to cartilagi-
nous glebae of greenish to brownish tint, except
Mesophelliaceae, which has a powdery gleba at
maturity.
Ecological diversity: most taxa form ecto-
mycorrhizae with various host trees, including
Pinaceae and Fagaceae in the Northern Hemi-
sphere and Myrtaceae (mostlyEucalyptusand
Leptospermum) and Nothofagaceae in the
Southern Hemisphere. In addition, some recent
studies have extended the range of hosts to
Caesalpiniaceae, Phyllanthaceae, and Diptero-
carpaceae (Castellano et al. 2000 ; Henkel et al.
2012 ). Phallogastraceae is the only family in the
order with a saprotrophic habit. Some species
ofHysterangiumform dense perennial hyphal
mats, which change the soil chemistry and
microorganism biomass (Griffiths et al. 1994 ).
Fruiting bodies of Hysterangiaceae and Meso-
phelliaceae are consumed by small mammals
and marsupials, and they often make up a sig-
nificant portion of the animals’ diet (Claridge
2002 ; Lehmkuhl et al. 2004 ).
Systematics: this order was proposed by
Zeller ( 1939 ), whose treatment was followed
by those of Locquin ( 1974 ) and Ju ̈lich ( 1981 ).
However, a Latin diagnosis was not provided
until Hosaka et al. ( 2006 ) formally described
the order. The order is strongly supported as
being monophyletic by multigene phylogenetic
studies (Hosaka et al. 2006 , 2008 ), but its rela-
tionships to other orders in Phallomycetidae
are not well supported.
Hysterangiales is divided into four families,
Hysterangiaceae, Mesophelliaceae, Gallacea-
ceae, and Phallogastraceae, all of which are
strongly supported as being monophyletic
(Hosaka et al. 2006 , 2008 ). Hysterangiaceae
and Mesophelliaceae form a clade, to which
Gallaceaceae is the sister family (Hosaka et al.
2006 ). Phallogastraceae, the only saprotrophic
member of the order, represents the earliest
branch, which is separated from the remaining
Hysterangiales (Hosaka et al. 2008 ), suggesting394 D.S. Hibbett et al.