Goniautoicous, occasionally dioicous. Plants erect, in loose or dense turfs or tufts. Stems simple or branched; central strand usually undifferentiated; axillary hairs usually sparse, short, with few cells, or abundant, elongate and multicellular. Leaves ±twisted and contorted when dry, the laminae generally inflexed to conduplicate, otherwise involute; moist leaves erect-spreading to reflexed, the laminae inflexed to plane; most leaves ±oblong or lanceolate-oblong; apex ±rounded, usually ±broadly obtuse or acute, otherwise narrowly acute, muticous, mucronate, apiculate or hair-pointed; margins mostly plane, or recurved, usually distally minutely (microscopically) crenulate, or minutely bluntly serrulate. Costa strong, ±prominently keeled on the abaxial surface, smooth to ±prorulose or papillose; adaxial surface (T.S.) with a single layer of laminal cells, 1–3 rows of cells with ±large lumina and strongly thickened walls; central strand distinct or undifferentiated, a small cluster of minute thin-walled cells; stereids in a single abaxial band, 2–8 cell rows with heavily thickened walls and minute lumina; abaxial epidermis ±undifferentiated. Distal medial laminal cells chlorophyllose, quadrate to subquadrate or short-oblong, their walls evenly thickened, usually bulging equally and papillose on both surfaces; papillae 2–8 over the lumina, ±C-shaped, or bulging more on the adaxial surface and bulging less and mammillose on the abaxial; distal marginal cells differentiated in 1 row, usually ovate, the narrow end projecting marginally, papillose or ±rhomboidal; upper corner projecting marginally; basal cells ±differentiated, ±rectangular, usually non-chlorophyllose, their transverse walls ±thickened, orange to yellow; longitudinal walls thin, hyaline to orange; surface walls usually smooth or papillose on the abaxial, as a group extending higher marginally or medially or not, or ±chlorophyllose, with the transverse and longitudinal walls thin or somewhat thickened, hyaline to pale orange or brownish; basal marginal cells undifferentiated or differentiated in a broad greenish band 2–20 narrow ±elongate cells. Specialised asexual reproduction usually lacking, alternatively as ±dense, filamentous, richly branched, dark brown brood bodies on stems.

Perichaetia terminal, the leaves ±sheathing or undifferentiated. Perigonia usually lateral, minute, bud-like, the leaves ±sheathing and paraphyses with undifferentiated distal cells; terminal perigonia with leaves differentiated or not, the paraphyses with enlarged distal cells. Calyptra persistent, mitrate, elongate-cylindrical, generally extending well below the capsule, smooth to ±prorulose or papillose, basally ±entire or fringed, distally narrowed to a ±elongate beak. Seta erect, short to elongate, straight to ±flexuose, ±twisted. Capsules stegocarpous, erect, the theca cylindrical, occasionally furrowed longitudinally or spirally; neck usually indistinct; annulus usually undifferentiated, massive, glossy, crimson-red, deciduous in large fragments; operculum conical, convex or concave-plane and short- to long-rostrate. Peristome absent or highly variable and with teeth in 1 or 2 concentric layers, ±lanceolate to ±linear and elongate. Spores highly variable in size, shape, polarity and ornamentation.

Encalyptaceae Schimp., Coroll. Bryol. Eur. 38 (‘1855’) [1856]. Type: Encalypta Hedw.

While the two genera of Encalyptaceae are differentiated solely by gametophytic features, remarkably there is convergence in sporophytic structure. This has led some to treat the monotypic Bryobrittonia as a species of Encalypta. Williams (1901) described Bryobrittonia on the basis of sterile plants, whereas Mitten (1864) had described fruiting plants as E. longipes Mitt. and erroneously reported the upper laminal cells to be papillose. This was a critical error, because the absence of papillae is one of the most distinctive features of Bryobrittonia. Nyholm (1998) noted that Bryobrittonia is indistinguishable from Encalypta sporophytically and concluded that Mitten’s treatment was correct. However, Steere (1953) felt that the distinctive gametophytic features “more than adequately” justify generic segregation, and Horton (1983) later concurred with his assessment. Bryobrittonia is restricted to tundra and montane boreal regions of North America and Eurasia.

Detailed developmental and structural analyses were the basis for Stone’s (1977) suggestion that a third genus, the monotypic Bryobartramia Sainsbury, might belong in the Encalyptaceae. However, she concluded that detailed studies of Pottiaceae, in particular, are necessary “before any valid assessment can be made”. Zander (1993) placed Bryobartramia in the Encalyptaceae based on shared characteristics of a large calyptra and large, coarsely papillose upper laminal cells that have a yellow KOH reaction. In contrast, Buck & Goffinet (2000) included only Encalypta and Bryobrittonia in the Encalyptaceae in the monotypic order Encalyptales and placed the monotypic Bryobartramiaceae in the Pottiales. The Encalyptaceae and Bryobartramia were among the outgroups included by Hedderson et al. (2004) in a phylogenetic analysis of haplolepideous mosses based on the chloroplast-encoded rps4 gene, and their data indicated that Bryobartramia is closely related to the Encalyptaceae. At the same time, Goffinet & Buck (2004) included Bryobartramia within the Encalyptaceae; however, more recently, Goffinet et al. (2008; 2012) treated Bryobartramia as a monotypic family in the Encalyptales.

The diversity of peristome structure among species of Encalypta, which Philibert (1884–90) interpreted as representing nematodontous and arthrodontous, diplolepideous and haplolepideous peristomes, led him to treat the Encalyptaceae as a basal group from which other mosses had diverged. Philibert suggested closest relationships with Polytrichaceae-Tetraphidaceae-Buxbaumia-Diphyscium, Orthotrichum and Pottioideae. Subsequently, the Encalyptaceae generally were placed close to Pottiaceae-Calymperaceae or Orthotrichaceae-Grimmiaceae until Edwards (1979; 1984) reported that the 2:3 pattern characteristic of haplolepideous mosses is not present in the Encalypta species he examined and that the Encalypta species Philibert considered nematodontous actually are arthrodontous. In 1984, Vitt proposed a new classification of Bryopsida in which he recognised two fundamental lineages of arthrodontous mosses with the link between the two being groups with diplolepideous peristomes in which the segments are opposite the teeth, including Funariineae, Buxbaumiineae and Encalyptineae. He placed the Encalyptineae just above the Buxbamiineae, the base of the otherwise haplolepideous lineage, and Funariineae at the base of the diplolepideous lineage. Subsequent studies have borne out Vitt’s (1984) recognition of close relationships among Encalyptaceae, Funariaceae and Buxbaumiaceae/Diphysciaceae, and a near-basal phylogenetic position of Encalyptaceae among Bryopsida. Newton et al.’s (2000) analyses of combined morphological and molecular data placed Encalypta and Bryobrittonia as a sister group to the haplolepideous mosses in an arthrodontous clade derived from three taxa (Diphyscium basal, with Funaria and then Timmia above) with diplolepideous, opposite peristomes. Analyses of nucleotide sequences from one nuclear and two chloroplast loci by Goffinet & Cox (2000) placed Encalyptineae as a sister group to Funariaceae in the lineage of arthrodontous mosses with opposite peristomes. Buck & Goffinet (2000) placed Encalyptales within the Funariidae beside Timmiales and Funariales. Later, Goffinet & Buck (2004) similarly included Encalyptales in the Funariidae, but next to Gigaspermales and Funariales and placed Timmiaceae in the Timiidae. They noted the general incongruence of gametophytic and sporophytic structure between Encalyptales and Funariales, but suggested that the latter might be most closely related to Encalyptales and the evidence for this could lie in development of the amphithecium. The Encalyptales are treated similarly by Goffinet et al. (2008, 2012). Further molecular support for the placement of Encalypta, Bryobrittonia and Bryobartramia in the Funariidae and Encalyptales is that these three taxa share with eight genera of Funariaceae and Discelium the 71-kb inversion in the large single copy of the plastid genome (Goffinet et al. 2007).

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