cation that resulted in the recognition of six
families (Humber 1989 ). Balazy’s ( 1993 ) five-
family classification of the Entomophthorales
follows the Batkoan classification more closely
than those developed later (Ben-Ze’ev and Ken-
neth 1982 ; Humber 1981 , 1989 ; Remaudie`re and
Keller 1980 ). The majority of the research on
Entomophthoromycota is on insect pathogens
because these taxa are the most common and
economically important members of the order.
Fewer mycologists have studied the saprobic
Entomophthoromycota, but they are also taxo-
nomically diverse. The genusAncylistes was
long treated as the type genus of an order of
zoosporic fungi until it was transferred to Ento-
mophthorales (Berdan 1938 ).
Partial zygomycete phylogenies based on
18S rDNA for Entomophthorales and Kickxel-
lales are available (Jensen et al. 1998 ; Nagahama
et al. 1995 ; O’Donnell et al. 1998 ). There is a
phylogenetic analysis using 18S and 28S rDNA
sequences from the medically important spe-
cies of Basidiobolales, Entomophthorales, Mor-
tierellales, and Mucorales (Voigt et al. 1999 ).
Two phylogenetic analyses, based on three
genes, include at least one member of every
recognized genus of Mucorales in culture
(O’Donnell et al. 2001 : 18S rRNA, 28S rRNA,
Ef-1a, morphology; Voigt and Wo ̈stemeyer
2001 : 18S rRNA, actin, Ef-1a). O’Donnell et al.
( 2001 ) revealed that only a few of the morpho-
logical characteristics, such as spore appen-
dages and trophocysts, were phylogenetically
informative.
A multigene phylogenetic study of all fun-
gal phyla, including zygomycotan fungi, was
published by Lutzoni et al. ( 2004 ); this data
set was composed of nucLSU and nucSSU
for Chytridiomycota, Glomeromycota, and
zygomycotan fungi, whereas additional
sequences were included for the remaining
phyla (Ascomycota, Basidiomycota).
Multigene phylogenies of the fungi, includ-
ing zygomycota, were subsequently published
(James et al. 2006 ; Liu et al. 2006 ). A cladogram
of the zygomycotan fungi based on three genes
(5.8S rRNA, 18S rRNA, 28S rRNA) was pub-
lished by White et al. (2006a). White ( 2006 )
presented a multigene phylogeny based on
rRNA from the available members of the Har-
pellales. Hibbett et al. ( 2007 ) published a new
classification of the fungi down to order based
on molecular phylogenetic studies of all fungal
groups, and four subphyla (Entomophthoro-
mycotina, Kickxellomycotina, Mucoromyco-
tina, Zoopagomycotina) with unknown
affinities were reported. Later, a fifth subphy-
lum, Mortierellomycotina, was described, also
with unknown affinities (Hoffmann et al. 2011 ).
Humber (2012b) described a new phylum,
Entomophthoromycota, and several higher taxa
for the entomophthoralean fungi. This classifi-
cation was based on a multigene phylogeny
presented by Gryganskyi et al. ( 2012 , 2013 ).
Two recent papers summarize the classification
of the zygomycotan fungi but differ in Mucor-
ales family recognition (Benny 2012 ; Voigt
2012 ).V. Reproduction and Dispersal
A. GrowthZygomyceous fungi can grow quite rapidly,
producing sporangiospores (or conidia) and,
under optimal conditions, even zygospores in
1–3 weeks. Many members of Mucorales and
Kickxellales are saprobes that can be encoun-
tered on diverse organic substrates. Many or all
taxa in Zoopagales, Dimargaritales, and Ento-
mophthoromycota are obligate parasites; a few
are nonhaustorial. Many members of Zoopa-
gales are mycoparasites, but the remaining
ones, and some members of Entomophthoro-
mycota, are parasites of nematodes or their
eggs, amoebae, rotifers, and tardigrades (Bar-
ron 2004 ; Tucker 1981 ). Other members of
Entomophthoromycota are obligate parasites
of algae (Ancylistes), fern prothallia (Comple-
toria), or insects or mites (most taxa); most
Basidiobolusand Conidiobolusspp. are sap-
robes (Humber 1989 ).B. DispersalSpore dispersal may be active (forcible dis-
charge), as inPilobolusand Entomophthoro-
mycota (except Massospora), or passiveZygomycetous Fungi: Phylum Entomophthoromycota and Subphyla Kickxellomycotina,... 215