Development of the enigmatic peristome of Timmia megapolitana (Timmiaceae; Bryophyta)

The Timmiaceae (Bryophyta) have been traditionally classified within the Bryales based on peristome architecture. Phylogenetic analyses of nucleotide sequences have revealed relationships that are incongruent with this hypothesis and have implicated an origin for this lineage early in the radiation of arthrodontous mosses (Bryopsida). This unexpected phylogenetic placement raises important questions about the evolutionary significance of the Timmia peristome, which differs from all other mosses by 64 isomorphic filaments topping the endostomial membrane. A developmental study of the peristome in Timmia megapolitana was undertaken to examine alignments of anticlinal cell walls in the inner peristomial layer (IPL) with those of the primary peristomial layer (PPL), a character that has been used to define major arthrodontous lineages. Criteria were established for assessing longitudinally homologous regions that contribute to the peristome-forming region. Young sporophytes were examined using histological techniques, and the alignment of the cell wall divisions was quantified. Critical divisions in the IPL of T. megapolitana were determined to be symmetrical, similar to patterns in the Funariales. This research provides novel developmental observations for a putative ancestral lineage of arthrodontous mosses, reevaluates criteria used to compare developmental studies of different lineages and discusses the phylogenetic implications of these observations.