One contribution of 15 to a theme issue 'Weird sex: the underappreciated diversity of sexual reproduction'. Haploid gametophytes of bryophytes spread by clonal growth but mate locally, within an area defined by the range of sperm movement. Rarity of establishment from spores or vegetative competition can result in unisexual populations unable to reproduce sexually. Females typically outcompete males, probably because females expend fewer resources than males on the production of gametes. Extreme sexual dimorphism-tiny males growing as epiphytes on much larger females-has evolved many times. Haploid selfing is common in bryophytes with bisexual gametophytes, and results in completely homozygous sporophytes. Spores from these sporophytes recapitulate the genotype of their single haploid parent. This process can be considered analogous to 'asexual' reproduction with 'sexual' reproduction occurring after rare outcrossing between haploid parents. Ferns also produce bisexual haploid gametophytes but, unlike bryophytes, haploid outcrossing predominates over haploid selfing. This difference is probably related to clonal growth and vegetative competition occurring in the haploid but not the diploid phase in bryophytes, but the reverse in ferns. Ferns are thereby subject to stronger inbreeding depression than bryophytes.This article is part of the themed issue 'Weird sex: the underappreciated diversity of sexual reproduction'.
Sexual lives of bryophytesAnimal life histories are characterized by multicellular diploid individuals that produce haploid gametes by meiosis. Gametes are the only haploid cells of such life cycles. The life histories of bryophytes (mosses, liverworts and hornworts) are fundamentally different. Multicellular haploid gametophytes (gameteproducing plants) produce gametes by mitosis. The fertilization of an ovum by a sperm produces a zygote that develops into a short-lived multicellular diploid sporophyte (spore-producing plant) that grows attached to the maternal gametophyte on which it depends for nutrients. Sporophytes produce haploid spores by meiosis that germinate to produce new gametophytes (figure 1). Mating is restricted to the distance over which sperm can find ova, usually in the millimetre to centimetre range [1]. Clonal expansion occurs via vegetative spread of gametophytes into neighbouring space and production of gemmae (asexual propagules) that disperse gametophytic clones to new sites [2].Gametophytes of dioecious bryophytes are unisexual, producing either eggs or sperm, but not both. Males and females compete for space but require close proximity for sexual reproduction. By contrast, gametophytes of monoecious bryophytes are bisexual, able to produce both eggs and sperm. A bisexual gametophyte that fertilizes its own eggs engenders homozygous sporophytes all of whose spores are genetically identical (excepting meiotic errors and postzygotic mutations). Roughly 70% of liverworts, 60% of mosses and 40% of hornworts are dioecious [3]. Bryophyte life cycles are not only fascinating in th...