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158<br />

James F. White Jr. et al.<br />

endophyte has not been found to possess insect deterrent properties, instead<br />

growth enhancements have been proposed (Faeth et al. 2000). Finally several<br />

species of fine fescues infected by Epichloë festucae were found to have an<br />

increased resistance to the dollar spot disease caused by Sclerotinia homeocarpa.<br />

It seems evident that <strong>plant</strong>s benefit tremendously from the colonization of<br />

symbiotic microbes. The benefits to hosting mutualistic microbes likely outweigh<br />

losses in terms of nutrient use by the microbes. The widespread level of<br />

infection of grasses by Epichloë and asexual forms in Neotyphodium are evidence<br />

that these associations also have evolutionary value. In one study of<br />

endophytes in grasses, infection levels in some hosts (e.g., Achnatherum<br />

robustum and Festuca versuta) were estimated to be greater than 90 % in populations<br />

throughout the ranges of the grasses (White 1987).<br />

3 Coevolution of Clavicipitalean Fungi with Grass Hosts<br />

It is evident that Epichloë (Clavicipitaceae; Ascomycetes) and related asexual<br />

endophytes have co-evolved with the cool-season (C-3) grasses in which they<br />

perennate (White 1988). Species of these endophytes are unknown from<br />

warm-season (C-4) grasses (White 1987). On the other hand, endophytic<br />

species in genus Balansia, also in family Clavicipitaceae, appear to have coevolved<br />

with warm-season grasses and are rarely or never found on cool-season<br />

hosts (White and Owens 1992). In co-evolving with grasses, it is logical to<br />

expect that their interactions with hosts became more sophisticated.<br />

4 The Jump from Insects to Plants<br />

4.1 Trans-Kingdom Jump<br />

Analysis of rDNA 26S sequence data indicates that the predominantly insectinfecting<br />

subfamily Cordycipitoideae (Clavicipitaceae) is the most deeply<br />

rooted group and is, therefore likely ancestral to grass-infecting species (Sullivan<br />

et al. 2000).A trans-kingdom host jump is postulated to have occurred to<br />

<strong>plant</strong>s. Such a jump could have occurred gradually through intermediate<br />

forms that were parasitic on both insects and <strong>plant</strong>s.<br />

4.2 Intermediate Stages in the Transition to Plants<br />

Several Cordycipitoideae exhibit stages of such a transition. Most of the<br />

Cordycipitoideae (e.g., Cordyceps militaris and C. sinensis) infect insect hosts<br />

and mummify them, using their necrotrophied bodies as energy to fuel fun-

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