environments, but it has a relatively high temperature
optimum for growth, so its use as a biological control
agent is restricted to glasshouse environments in the
cooler parts of the world. In addition to these conidial
fungi, several species of Zygomycota commonly cause
insect diseases in natural environments and also in field
crops. These fungi belong to a subgroup of Zygomycota
termed the Entomophthorales(see Fig. 2.11). They
include genera such as Entomophthoraand Pandora(pre-
viously called Erynia). They produce large sporangia at
the tips of their hyphae, and the sporangia are released
intact, functioning as dispersal spores (Fig. 15.1d). The
Entomophthorales often cause natural epizootics in
humid conditions, and attempts are being made to
develop them as commercial biocontrol agents.
The infection cycle
The general infection cycle of insect-pathogenic fungi
is summarized in Fig. 15.3. In almost all cases these fungi
initiate infection from spores that land on, and adhere
to, the insect cuticle. If the relative humidity is high
enough, the spore then germinates and usually forms
an appressorium, equivalent to the appressoria produced
by many plant-pathogenic fungi (see Fig. 5.2). The
appressorium often develops over an intersegmental
region of the cuticle, and in one of the common insect
pathogens, Metarhizium anisopliae, this behavior closely
parallels the behavior of the rust fungi (Chapter 5),
because the germ-tube recognizes the host surface
topography, or artificial surfaces with appropriate
FUNGAL PARASITES OF INSECTS AND NEMATODES 311
Fig. 15.2Beauveria bassianaand ex-
amples of the diseases that it causes.
(a) Spore-bearing structures of Beauveria
bassiana in laboratory culture. The
conidia typically develop in zigzag-like
chains on long conidiophores. (b) An
adult cicada beetle densely covered
with white sporulating pustules of B.
bassianathat have emerged through the
intersegmental plates of the insect
cuticle. (c) Sporulation of Beauveria
from between the cuticular plates of a
naturally infected green cockchafer
beetle. (d) Heavy infection of an adult
pecan weevil by Beauveria.(e) Grubs
of pecan weevil at different stages of
infection by Beauveria; healthy grubs are
shown at left. ((a),(b) Courtesy of G.L.
Barron. (c) Courtesy of Shirley Kerr;
http://www.kaimaibush.co.nz/. (d),(e)
Courtesy of Louis Tedders (photogra-
pher) and USDA, Agricultural Research
Service.)