to infect other conifers and broad leaved trees as well. In contrast, H. parviporum lives almost exclusively on Picea abies, but attacks Abies sibirica in northeastern Europe 11,12 , while H. abietinum is a pathogen or saprophyte on Abies species in southern and central Europe 11,12. The North American species H. irregulare mostly infects Pinus and Juniperus species but also Abies balsamea while H. occidentale has a host range excluding Pinus spp. but including species in the genera Abies, Picea, Pseudotsuga, Tsuga, Sequoiadendron 13. Although there is a patter in the host preferences among the H. annosum s.l. species the specialization has not been driven by co-speciation together with the host 10. The speciation of H. annosum s.l. complex has been suggested to start with a split between an ancestor of the pine infecting species H. annosum s.s. and H. irregulare and the ancestor of the non-pine infecting species H. parviporum, H. abietinum and H. occidentale in Lauraisa 10. The common ancestor of H. annosum s.s./H. irregulare migrated from west Europe to eastern North America. After the Atlantic land bridge disappeared the two species H. annosum s.s. and H. irregulare evolved in allopatry for several millions of years in Europe and North America, respectively. While the ancestor of H. parviporum, H. abietinum and H. occidentale may originate in east Asia or west North America 10. In North America the H. irregulare continued to spread west over the continent while the spread of H. occidentale was restricted to the east by an arid region which probably was colonized by pines, which are a nonhost for H. occidentale 10. After original speciation between pine and non-pine infecting species, H. irregulare and H. occidentale have had a long period to evolve in allopatry before today's overlapping geographic distribution in the western North America. How the evolutionary history with speciation in sympatry and a long period of allopatric evolution have shaped their genomes and genetic differences found between H. irregulare and H. occidentale is not well understood. Comprehensive studies of H. irregulare have revealed a trade off in gene expression between saprotrophic and parasitic lifestyle 14. During the saprotrophic growth H. irregulare use various cellulose and pectin degrading enzymes and many cellulose oxidoreductase genes are up-regulated during wood degradation while genes related to secondary metabolism are more commonly activated during growth in bark 14. Transcripts identified from H. annosum s.s. and Norway spruce interaction showed similar gene induction patterns as seen in H. irregulare 15. Furthermore, a global gene expression study of H. annosum s.s. found that the glyoxylate cycle might be important for fungal adaptation to change of nutrients availability in the environment, melanization for adapting to salt stress, and the pathway related to DNA repair for oxidative stress 16. We hypothesize that H. irregulare and H. occidentale evolved distinct genetic machineries for infection and colonization of t...