The genera that constitute the Pucciniastreae display a wide variation in the type of teliospore produced, as well as in the time and place of production. From the standpoint of development, however, there is a general situation that is common. In all genera primordial cells are formed from enlarged hyphal cells of the mycelium. These give rise to teliospore initials which are in the epidermal cells in Calyptospora, Milesia, Hyalopsora and Thecopsora, and are subepidermal in the other genera. These initials divide to form the mature teliospores, which are thick or thin walled, and few to many-celled. In all cases the teliospore is the product of a single primordial cell.The teliospores of Calyptospora goeppertiana are formed from a perennial mycelium, which causes a witches' broom and hypertrophied stems on species of Vaccinium. The mycelium gives rise to primordial cells in the cortex just below the epidermis. Each primordial cell pierces the host wall above and the contents pass in to form the initial, which by growth and division becomes the teliospore. The mature teliospores are one- to four-celled, with a thickened, dark brown wall. Development is not simultaneous, but progressive, and the teliospores are first formed in the basal parts, moving slowly upward until every cell of the hypertrophied portion of the stem is completely filled. In four species of Milesia the method of development is similar. The spores, however, are thin walled, and are formed in the epidermal cells of the overwintered fronds of their fern hosts. Thecopsora vacciniorum is similar to Milesia in many respects. The teliospores are intra-epidermal, thin walled and multicellular. In Pucciniastrum the teliospores are subepidermal, and arise from primordial cells, as in Calyptospora, Milesia and Thecopsora. The teliospore initials are closely packed, and the mature spores may form extended crusts. The simplest type of development is found in Uredinopsis, which is generally considered to be the most primitive of the fern rusts. Primordial cells are formed in the same way as in the other genera. These round up to form the initials, and cross walls are laid down to give the mature spores.From these studies two possible lines of development are suggested, both beginning with Uredinopsis. One line would lead through the intra-epidermal forms, as Milesia, Calyptospora, etc., and the other through the subepidermal genera, as Pucciniastrum and Melampsoridium.
Unh1, an Ustilago maydis Ndt80-like protein, controls completion of tumor maturation, teliospore development, and meiosis