This review provides a synopsis of apogamous reproduction in ferns and highlights important progress made in the study of fern apomixis over the past decade. First, a summary of the apomictic fern life cycle is provided, distinguishing between two pathways to diploid spore production that have been documented in apomictic ferns (premeiotic endomitosis and meiotic first division restitution) and briefly discussing the evolutionary implications of each. Next, recent trends in fern apomixis research are highlighted, exposing a shift in focus from the observation and characterization of apomixis in ferns to more integrated studies of the evolutionary and ecological implications of this reproductive mode. Peer-reviewed contributions from the past decade are then summarized, spanning the identification of new apomictic lineages through to the developmental, phylogenetic, and population genetic insights that have been made in studies of fern apomixis during that time. Gaps in our understanding of fern apomixis are also discussed, including the extent and implications of recombinant apomixis in ferns, the possible reversibility of reproductive mode (from apomictic to sexual) in ferns, and the genomic causes and consequences of apomixis in seed A u t h o r M a n u s c r i p t A u t h o r M a n u s c r i p t A u t h o r M a n u s c r i p tGrusz, 2 of 23 free vascular plants. Finally, future directions for fern apomixis research are discussed in the context of modern technological advances and recent insights from studies of apomixis in other groups.Keywords: apogamy, evolution of sex, meiosis, pteridophytes.A u t h o r M a n u s c r i p t A u t h o r M a n u s c r i p t A u t h o r M a n u s c r i p t
Grusz, 3 of 23 "…apomixis is an escape from sterility, but it is an escape into a blind alley of evolution…" -Darlington, 1939A renaissance is underway in fern biology, propelled, in part, by technological advances being embraced by a growing research community. In particular, modern, cost-effective tools are enabling new inquiries into the unique fern life cycle, which alternates between independent, free-living sporophyte and gametophyte generations (Haufler et al., 2016). Among the many aspects of fern reproduction that are being examined in light of new approaches-and an improved understanding of fern evolution as a whole-is their propensity for apogamous reproduction (i.e., apomixis). This form of asexual reproduction (by spore or seed) is more common in ferns than in any other group of vascular plants. Nearly 10% of the ferns for which reproductive mode has been determined exhibit apomixis (Walker, 1985; Liu et al., 2012), compared to less than 1% in flowering plants (Bicknell & Koltunow, 2004; Becks & Alavi, 2015). Given that data on reproductive mode is lacking for the vast majority of ferns, this relative contribution of apomicts to fern diversity may well be an underestimate (Liu et al., 2012). The elevated frequency and distinctive meiotic characteristics of apomixis in ferns, which are elaborated upon bel...