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Insect Control: Biological and Synthetic Agents - Index of

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390 11: Entomopathogenic Fungi <strong>and</strong> their Role in Regulation <strong>of</strong> <strong>Insect</strong> Populations<br />

(Jensen et al., 1998), <strong>and</strong> some groups, such as<br />

the Glomales, may be placed elsewhere eventually<br />

(e.g., Bruns et al., 1993). Basidiobolus ranarum<br />

<strong>and</strong> other Basidiobolus species, traditionally considered<br />

as Zygomycetes, have now been placed in<br />

the Chytridiomycetes, despite being nonflagellate<br />

(Nagahama et al., 1995).<br />

Figure 2 Tick (Ixodes ricinus) killed by Metarhizium anisopliae.<br />

(Photo courtesy <strong>of</strong> C. Nielsen.)<br />

Within the true Zygomycota, the class Trichomycetes<br />

contains species <strong>of</strong>ten associated with insects.<br />

While Sweeney (1981) described the species Smittium<br />

morbosum (Trichomycetes) as a pathogen <strong>of</strong><br />

mosquitoes, most associations <strong>of</strong> the Trichomycetes<br />

are symbiotic or weakly parasitic rather than true<br />

pathogens (Beard <strong>and</strong> Adler, 2002; Cafaro, 2002).<br />

Some species <strong>of</strong> Mucor (Mucorales) are occasionally<br />

associated with insect mortality.<br />

The majority <strong>of</strong> entomopathogenic species within<br />

Zygomycota are contained in one order, the Entomophthorales.<br />

More than 200 entomopathogenic<br />

Entomophthorales species have been recognized.<br />

They commonly cause spectacular epizootics <strong>and</strong><br />

are characterized in all genera but one (Massospora),<br />

by the production <strong>of</strong> forcibly discharged<br />

primary conidia. Many species are capable <strong>of</strong> producing<br />

various types <strong>of</strong> secondary conidia from the<br />

primary conidia <strong>and</strong>, in some cases, infection is<br />

obligatorily through a secondary conidium. Many<br />

species are also capable <strong>of</strong> producing resting spores,<br />

long-lived zygospores or azygospores. These fungi<br />

are generally obligate pathogens in nature, <strong>and</strong><br />

many species are presently difficult or impossible<br />

to culture on artificial media.<br />

Until around the 1960s, most entomopathogenic<br />

Entomophthorales were contained within a<br />

single genus, Entomophthora, but several years <strong>of</strong><br />

taxonomic revision have placed the species into<br />

several genera. The main genera containing<br />

entomopathogenic species are Conidiobolus, Entomophaga,<br />

Entomophthora, Erynia, Furia, Massospora,<br />

Neozygites, P<strong>and</strong>ora, Strongwellsea, <strong>and</strong><br />

Zoophthora.<br />

Figure 3 Adult cabbage fly (Delia radicum) killed by Beauveria bassiana. This fungus produces clusters <strong>of</strong> white conidia. (Photo<br />

courtesy <strong>of</strong> J. Eilenberg.)

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