Molecular and pathogenic variation within Melampsora on Salix in western North America reveals numerous cryptic species

In North America Melampsora rusts that parasitize willows (Salix species) have never been adequately studied and mostly have been referred to a collective species, Melampsora epitea (Kunze & Schm.) Thum, of European origin. Even taxa that are nominally distinct from M. epitea, such as M. abieticaprearum and M. paradoxa, currently are considered to be "races" of M. epitea. Within the range of our field surveys and collections in the Pacific Northwest and the Southwest only two species of Melampsora thus were expected: M. epitea (including its races) and M. ribesii-purpureae. In this study of Melampsora on 19 species of Salix in the western United States 14 phylogenetic species, or phylotypes, were apparent from nuclear rDNA sequencing of 140 collections or isolates. Our collections of the races of M. epitea, M. abieti-caprearum and M. epitea f. sp. tsugae belonged to one phylotype, termed lineage 'N'. Assuming that M. ribesii-pulpureae represents one other phylotype, 12 phylotypes still are unaccounted for by current taxonomy. Moreover Eurasian M. ribesii-purpureae was not closely related to any of the phylotypes reported here. Even more problematic was the resistance of Eurasian species of Salix, including the type host of M. epitea, S. alba, to North American Melampsora, including phylotype 'N', in both the field and in inoculation experiments. These results suggest the need for the description of many new species of Melampsora on Salix in western North America. Additional analyses presented here might guide further research in this direction.