Phytopathogens often establish and proliferate on specific plant organs and tissues, triggering leaf spots, stem cankers, fruit or root rots, and tuber scabs. A pathogen's structural specificity is often caused by adaptation to a given organ (leaf, stem, flower, root, etc.), tissue (xylem, phloem, mesophyll, etc.), or a developmental stage of the host (Barrett and Heil, 2012). Such specificity can be expressed at the level of its entry tissue, its colonized tissue, its sporulating tissue, or any combination of these factors. Pathogens can exert high structural specificity, such as powdery mildew species confined to the leaf epidermis, or have no structural specificity, such as Sclerotinia sclerotiorum, a generalist capable of infecting virtually any tissue. Furthermore, a pathogen's structural specificity can change throughout its lifecycle (e.g., Figure 1g,h). Both structural specialists and generalists may have benefits considering infection strategies. Examples of such advantages include performance, competition, or niche expansion (Barrett and Heil, 2012). However, research on the processes driving structural specificity is still in its infancy and many questions are only partially