Premise of research. The Phosphatidylethanolamine-Binding Protein (PEBP) transcription factors are critical components of the flowering pathway. However, they are present in all vascular plants, even in those lacking flowers. It is unclear whether the same factors were already recruited in nonflowering plant reproductive transitions often linked to sporogenesis and leaf dimorphism. Here, we provide an inclusive phylogenetic framework for PEBP genes and study their spatiotemporal expression patterns in selected lycophytes and ferns.Methodology. We generated reference transcriptomes for four fern species, Adiantum raddianum, Anemia villosa, Equisetum bogotense, and Equisetum giganteum. Maximum likelihood analysis of PEBP genes was performed across land plants. Expression analysis by reverse transcriptase PCR and in situ hybridization analysis were carried out for the lycophyte Selaginella moellendorffii and the fern A. raddianum.Pivotal results. We (1) confirm that the FLOWERING LOCUS T (FT) flowering activator gene and TERMINAL FLOWER 1 (TFL1) flowering transition repressor gene are seed plant specific, (2) identify that ferns possess FT/TFL1 genes, which predate the duplication resulting in the FT and TFL1 clades, and (3) detect that lycophytes and ferns have MOTHER OF FT AND TFL1-like (MFT) genes, which are redundant in flowering activation in model angiosperms. Expression analyses show that the lycophyte and fern PEBP genes are broadly expressed in vegetative and reproductive stages and that most species-specific copies overlap during sporangium initiation and development.Conclusions. The data point to a redundancy of PEBP copies in the reproductive transitions and spore-producing stages and suggest that PEBP genes are involved in meristem maintenance and sporangium development in both lycophytes and ferns. In turn, it is possible that PEBP genes have putative plesiomorphic roles in reproductive transitions across tracheophytes.