The nature and distribution of vegetative desiccation-tolerance in hornworts, liverworts and mosses

Desiccation-tolerance is the unique ability to revive from the air-dried state. In this review, I summarize the distribution of vegetative desiccation-tolerance within the land plants emphasizing bryophytes, and provide a current checklist of those moss, liverwort and hornwort species with documented and experimentally determined tolerance. Desiccation-tolerant species can survive equilibration with either modestly dry air (i.e., 70–80% RH or −30 to −48 MPa) or extremely dry air (i.e., 0–30% RH or less than −162 MPa), and have been identified within seven classes of bryophytes, the Andreaeopsida, Bryopsida, Polytrichopsida and Tetraphidopsida (mosses), Jungermanniopsida and Marchantiopsida (liverworts) and the Anthocerotopsida (hornworts); 210 out of 21,000 bryophyte species (ca. 1.0%) have been experimentally determined to possess vegetative desiccation tolerance—158 species of mosses, 51 species of liverworts and one species of hornwort. Finally, I propose a comprehensive survey of mosses, liverworts and hornworts using a standard procedure that employs modulated chlorophyll fluorescence for analyzing vegetative desiccation-tolerance (i.e., the “Austin Protocol”).