Cryo-scanning electron micrographs of freeze-fractured hornwort gametophytes (a–c) and sporophytes (d–i): Anthoceros agrestis (a,c,d–f); Folioceros fusiformis (b); Leiosporoceros dussii (g); Megaceros enigmaticus (h); Dendroceros granulatus (i). Sections through thalli showing mucilage-filled cavities (asterisk). (c) Nostoc colony. (d,g) Intercellular spaces are initially liquid-filled (asterisk) but become gas-filled (e, arrowed) following stomatal opening. (f) Columella with gas-filled (asterisk) intercellular spaces. (h,i) Young (h) and mature (i) sporophytes of astomate taxa, showing complete absence of intercellular spaces in the assimilatory layers which collapse and dry (i). Scale bars: (a,b) 200 µm; (d,e,g) 50 µm; (c,f,h,i) 20 µm.