as mushroom parasites.DicranophoraandSpi-
nellusare psychrophilic and only grow at tem-
peratures below 20 C; 15 C is optimal
(Voglmayr and Krisai-Greilhuber 1996 ; Watson
1964 ).Syzygitescan grow at 25C but not 30C
(Wenger and Lilly 1966 ). All three genera grow
in pure culture on nutrient-rich media.Syzy-
gites megalocarpusEhrenb.: Fr. has the most
extensive host range (98 species in 22 basidio-
mycete families and 4 ascomycete species)
(Kovacs and Sundberg 1996 ).
The facultative parasites of mushrooms,
arthropods (Sporodiniella umbellata), and
other members of Mucorales (Lentamyces par-
ricida,Chaetocladiumspp.,Parasitella simplex)
may only be found on a few, or many, hosts in
nature and can all be grown on ordinary culture
media. Those parasitic taxa that grow axeni-
cally on culture are usually not altered morpho-
logically. The mucoralean mycoparasites
Chaetocladium, Lentamyces, and Parasitella
are nonhaustorial biotrophic fusion parasites
that induce gall formation in a host (Hoffmann
and Voigt 2009 ).Sporodiniellaonly is found in
nature on insects and spiders in the subtropics
and tropics (Chien and Hwang 1997 ; Evans and
Samson 1977 ).
- Zoopagomycotina and Zoopagales
All members of Zoopagales sensu Benjamin
( 1979 ) (Fig.8.5) are obligate parasites (preda-
ceous, ectoparasites, or endoparasites) of
other fungi or small animals (amoebae, roti-
fers and nematodes and their eggs), with pre-
daceous species and ectoparasites forming
haustoria in the host. The hyphae are coeno-
cytic or septate. The thallus is a branched or
unbranched, inflated hyphal coil in the host, or
it consists of branched external hyphae. Asex-
ual reproduction is by conidia [Fig.8.5(42)] or,
possibly, unispored [Fig.8.5(43)] (G.L. Benny,
unpublished data) or multispored merospor-
angia [Fig.8.5(44, 47)]. The conidia are formed
as single spores or in simple or branched
chains of many merosporangia. Other means
of asexual reproduction are by arthrospores
and chlamydospores. Sexual reproduction is
by the formation of zygospores[Fig.8.5(45,
46, 48)]. Zygospores, when formed, are more
or less globose, have opposed suspensors, and
either do not [Fig.8.5(45)] or may bear suspen-
sor outgrowths [Fig.8.5(46, 48)].
Zoopagales contained only the type family
until Duddington ( 1973 ) removed the ecto- and
endoparasites and transferred them to Cochlo-
nemataceae, leaving only the predaceous forms
in the Zoopagaceae. Benjamin ( 1979 ) trans-
ferred the haustorial parasitic members (Heli-
cocephalidaceae, Piptocephalidaceae) of
Mucorales sensu Hesseltine and Ellis ( 1973 )to
Zoopagales. Helicocephalidaceae, including
Rhopalomyces, are parasites of nematodes and
their eggs and of rotifer eggs, whereas Piptoce-
phalidaceae are mycoparasites, especially of
Mucorales. The mycoparasitic members of Zoo-
pagales are restricted to the Piptocephalidaceae
and Sigmoideomycetaceae.Piptocephalis and
Syncephalis (Piptocephalidaceae) (Benjamin
1959 ) contain numerous species and may be
found anywhere the host (usually a member of
the Mucorales) occurs. These genera are more
or less cosmopolitan, whereas the third genus,
Kuzuhaea (Benjamin 1985a), is known only
from the original isolation in Japan.
Ultrastructural evidence on the number
and appearance of wall layers shows that the
asexual spores of Stylopage rhabdospora
Drechsler (Saikawa 1986 ) are catenate conidia
with a wall structure like that of Entomophthor-
ales (Benny et al. 2001 ) [Fig.8.4(32–35)] rather
than merosporangiospores, as suggested by
Benjamin ( 1979 ). The other members of
Cochlonemataceae and Zoopagaceae that have
been examined ultrastructurally also produce
conidia (Saikawa2011b).
Saikawa and coworkers made several
observations on selected taxa of two families
of Zoopagales (Cochlonemataceae, Zoopaga-
ceae) using both light and electron microscopy.
Saikawa et al. ( 2011 ) described the ontogeny
and germination of Acaulopage pectospora
zygospores. Observations made by Saikawa
and Morikawa ( 1985 ) and Saikawa et al.
( 1988 ) indicated thatA. pectosporawas similar
toZoophagus insidiansSommerst. Dick ( 1990 )
observed the zygospores of A. pectospora,
transferred the species to Zoophagus, and
revised the genus description. A phylogeneticZygomycetous Fungi: Phylum Entomophthoromycota and Subphyla Kickxellomycotina,... 235