Plants medium-sized to large, sometimes slender and wiry. Stolons creeping; rhizoids from clus-ters of initials abaxial to the leaf insertions, on stolons, and at the base of primary stems. Stems erect to pendent, typically stipitate, regularly pinnately to bipinnately branched; branches determinate or indeterminate, evenly foliate; stems in cross section with a well-developed sclerodermis (sometimes with a unistratose hyalodermis), central strand absent; paraphyllia absent; pseudoparaphyllia nar-rowly foliose to filamentous; axillary hairs 2-celled. Stipe leaves appressed or appressed below and squarrose above; margins often narrowly recurved. Stem and branch leaves smooth or undulate to ru-gose, decurrent; margins subentire, serrulate, or serrate, plane, recurved or inflexed, at times bordered by elongate to sigmoid cells; costae strong, double, reaching above midleaf, often subpercurrent, par-allel or diverging throughout, at times with a dorsal crest, usually ending in a stout tooth, in cross sec-tion with homogeneous, thick-walled cells; cells short, thick-walled, smooth or prorate, occasionally porose; alar cells not or weakly differentiated. Asexual propagula often present on costae or leaf mar-gins. Dioicous, synoicous, or autoicous. Setae short, flexuose, smooth or roughened above. Capsules exserted or immersed, erect, oblong-cylindric, symmetric; exothecial cells thick-walled; stomata usu-ally present; opercula short-rostrate; annuli rudimentary; peristome diplolepideous; exostome teeth pale yellow, narrowly triangular, dorsal (outer) surface papillose throughout, trabeculae and median line faint; endostome yellow, lightly papillose throughout, basal membranes low, segments not or nar-rowly perforate, not keeled, cilia rudimentary or absent. Calyptrae mitrate, with archegonia and/or sparse hairs.