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Several species of the insect pathogenic fungus  Metarhizium  are associated with certain plant types and genome analyses suggested a bifunctional lifestyle; as an insect pathogen and as a plant symbiont. Here we wanted to explore whether there was more variation in genes devoted to plant association ( Mad2 ) or to insect association ( Mad1 ) overall in the genus  Metarhizium . Greater divergence within the genus  Metarhizium  in one of these genes may provide evidence for whether host insect or plant is a driving force in adaptation and evolution in the genus  Metarhizium . We compared differences in variation in the insect adhesin gene,  Mad1 , which enables attachment to insect cuticle, and the plant adhesin gene,  Mad2 , which enables attachment to plants. Overall variation for the  Mad1  promoter region (7.1%),  Mad1  open reading frame (6.7%), and  Mad2  open reading frame (7.4%) were similar, while it was higher in the  Mad2  promoter region (9.9%). Analysis of the transcriptional elements within the  Mad2  promoter region revealed variable STRE, PDS, degenerative TATA box, and TATA box-like regions, while this level of variation was not found for  Mad1 . Sequences were also phylogenetically compared to EF-1α, which is used for species identification, in 14 isolates representing 7 different species in the genus  Metarhizium . Phylogenetic analysis demonstrated that the  Mad2  phylogeny is more congruent with 5′ EF-1α than  Mad1 . This would suggest that  Mad2  has diverged among  Metarhizium  lineages, contributing to clade- and species-specific variation, while it appears that  Mad1  has been largely conserved. While other abiotic and biotic factors cannot be excluded in contributing to divergence, these results suggest that plant relationships, rather than insect host, have been a major driving factor in the divergence of the genus  Metarhizium .

Variability in the Insect and Plant Adhesins, Mad1 and Mad2, within the Fungal Genus Metarhizium Suggest Plant Adaptation as an Evolutionary Force