The erect secondary thallus (podetium) of the lichen Cladonia grayi de
velops in distinct stages that can be traced to the development of its
fungal meristem tissue. In early ontogeny, while the contiguous meris
tem grows unilaterally both outward and vertically, the gross morpholo
gy of the podetium resembles other cup-forming species. In later ontog
eny, when the divided meristem grows vertically, the podetium forms a
subtly articulated spiral. The spiral growth form is most apparent in
branching proliferations that grow along the cup margin. The meristem
tissue at the apex of these proliferations is curved, maturing sequent
ially in a spiral pattern. The proliferations curve in a spiral around
the vertical axis of the podetium. Apothecia at the apex of prolifera
tions curve in a similar manner. Evidence for spiral growth along the
cup margin can also be inferred from the sequential development of pyc
nidia that emerge from it. Similar to early land plants, the growth dy
namics of C. grayi may represent an evolutionary advance from an ortho
tropous growth pattern to a branched, photosynthetic thallus.