Bryaceae
Dioicous, synoicous or, rarely autoicous (not in Victoria). Asexual reproduction by tubers on rhizoids or stems, by gemmae or bulbils on stems or in leaf axils, or by leafless terminal shoots (not in Victoria) or deciduous leaf axil brood branchlets (not in Victoria). Tufts, turves, cushions or scattered individuals on soil, rocks, logs or trees. Stems erect, sometimes arising from stoloniferous primary stems, simple, forked, or branched by innovation below sporophytes, with coloured rhizoids toward base; central strand present. Leaves arranged around stem and facing all directions or rarely complanate, often larger and more crowded toward stem apex, frequently in comal tufts or rosulate, erect and imbricate to wide-spreading when moist, scarcely altered, or twisted or crisped when dry; apex broadly rounded, acute or acuminate, often with a hairpoint; costa subpercurrent to excurrent; margin entire to serrate, plane, recurved, or revolute, frequently with a border of more elongated cells; laminal cells rhomboidal or hexagonal to linear, similar throughout, or gradually becoming more elongate and rectangular toward leaf base or abruptly becoming quadrate to short-rectangular and often broader toward base, smooth; alar cells not or poorly differentiated. Acrocarpous. Capsule mostly inclined to pendent, rarely erect, usually symmetric, ovoid, pyriform or oblong-cylindric, rarely subglobose (not in Victoria), exserted, operculate; annulus usually present. Calyptra cucullate, smooth, glabrous. Operculum convex to short-conic, sometimes umbonate or apiculate, rarely short-rostrate. Peristome double and alternate, or rarely single (not in Victoria) or absent (not in Victoria); exostome of 16 entire teeth or rarely absent (not in Victoria); endostome of 16 segments arising from a generally well-developed basal membrane, or segments absent (not in Victoria) or fused with exostome (not in Victoria); cilia present or absent.
A cosmopolitan family comprising around 15 genera and 600 species; five genera and 36 species in Victoria.
The traditional circumscription of the Bryaceae included the Victorian genera Leptobryum, Pohlia, Schizymenium, and Orthodontium (Brotherus 1925). Phylogenies of chloroplast and nuclear DNA sequences have shown that these genera belong to separate lineages, closer related to other long-recognised families (e.g. Cox & Hedderson 1999; Cox et al. 2000). Leptobryum is closely related to Meesia and is placed in the Meesiaceae, Pohlia and Schizymenium are closely related to Mnium and now generally included in the Mniaceae, Orthodontium is closer related to the pleurocarpous mosses and is recognised in its own family, and the subfamily Bryoideae (Brotherus 1925), along with a few additional misplaced species and with minor adjustments (Cox & Hedderson 2003), comprises a single lineage now generally accepted as the Bryaceae (e.g. Buck & Goffinet 2000). Most species in the Bryaceae have a combination of broad, hexagonal or rhomboid-hexagonal laminal cells and pendent elongate pyriform capsules with a long neck and well-developed double-alternate peristomes (Cox & Hedderson 2003; Spence & Ramsay 2006), but features shared by all members of the family are lacking (Pederson et al. 2003).
Generic delimitation within the family is highly problematic. Several genera have been described from a broadly defined Bryum based on morphology (Spence 1996, 2005; Spence & Ramsay 2005). These do not correspond to well-supported lineages in phylogenies of DNA sequences from all genomic compartments (Cox & Hedderson 2003; Pedersen & Hedenäs 2003; Pedersen et al. 2003; Holyoak & Pedersen 2007; Pedersen et al. 2007). However, the traditional broad concept of Bryum and other traditionally recognised genera in the Bryaceae, which were largely based on sporophytic characters, are also non-monophyletic (Cox & Hedderson 2003; Pederson et al. 2003; Wang & Zhao 2009). These issues have been partly resolved by recognising some of the lineages obtained in molecular phylogenies as new genera or by revising previously described genera to better match lineages in molecular phylogenies (Pedersen 2005; Pedersen & Hedenäs 2005). However, several of these lineages are ill-defined morphologically (Pedersen et al. 2003; Pedersen & Hedenäs 2005; Holyoak & Pedersen 2007; Pedersen et al. 2007). This is particularly problematic for contemporary implementation of classification because few species have been included in phylogenetic analyses and their placement into a genus is met with uncertainty due to poorly known morphological limits or lack thereof for current putatively monophyletic genera. These uncertainties will likely persist until further sampling of taxa and DNA regions can better establish the boundaries of monophyletic genera within the Bryaceae.
The generic placement of species adopted here mainly follows the classification of Spence & Ramsay (2006) previously used for Australian taxa. Exceptions to this classification are made for genera that are markedly non-monophyletic or for species that have been shown by DNA phylogenies to be misplaced in the Spence & Ramsay (2006) classification. Consequently, Gemmabryum, which is markedly polyphyletic and has a type (G. pachythecum (Müll.Hal.) J.R.Spence & H.P.Ramsay) closely related to the type of Bryum (Pederson et al. 2003) is not accepted and Rosulabryum capillare (Hedw.) J.R.Spence is placed in Ptychostomum based on its closer relatedness to this genus (Holyoak & Pedersen 2007).
SpinningBrotherus, V.F. (1925). Musci (Laubmoos), in Engler, A. (ed.), Die natürlichen Pflanzenfamilien, edition 2. Bd 11. Engelmann, Leipzig.
Buck, W.R.; Goffinet, B. (2000). Morphology and classification of mosses, in Shaw, A.J. & Goffinet, B. (eds.), Bryophyte Biology, pp. 71–123. Cambridge University Press, Cambridge.
Cox, C.J.; Goffinet, B.; Newton, A.E.; Shaw, A.J.; Hedderson, T.A.G (2000). Phylogenetic relationships among the diplolepideous-alternate mosses (Bryidae) inferred from nuclear and chloroplast DNA sequences. The Bryologist 103: 224–241.
Cox, C.J.; Hedderson, T.A.J. (1999). Phylogenetic relationships among the ciliate arthrodontous mosses: evidence from chloroplast and nuclear DNA sequences. *Plant Systematics and Evolution * 215: 119–139.
Cox, C.J.; Hedderson, T.A.J. (2003). Phylogenetic relationships within the moss family Bryaceae based on chloroplast DNA evidence. Journal of Bryology 25: 31–40.
Holyoak, D.T.; Pedersen, N. (2007). Conflicting molecular and morphological evidence of evolution within the Bryaceae (Bryopsida) and its implications for generic taxonomy. Journal of Bryology 29: 111–124.
Pedersen, N. (2005). Validation of Imbribryum (Bryaceae). The Bryologist 108: 449.
Pedersen, N.; Cox, C.J.; Hedenäs, L. (2003). Phylogeny of the moss family Bryaceae inferred from chloroplast DNA sequences and morphology. Systematic Botany 28: 471–482.
Pedersen, N.; Hedenäs, L. (2003). Phylogenetic investigations of a well supported clade within the acrocarpous moss family Bryaceae: evidence from seven chloroplast DNA sequences and morphology. * Plant Systematics and Evolution* 240: 115–132.
Pedersen, N.; Hedenäs, L. (2005). Taxonomic and nomenclatural implications of phylogenetic studies of the Bryaceae based on molecular data and morphology. The Bryologist 108: 123–128.
Pedersen, N., Holyoak D.T.; Newton, A.E. (2007). Systematics and morphological evolution within the moss family Bryaceae: A comparison between parsimony and Bayesian methods for reconstruction of ancestral character states. Molecular Phylogenetics and Evolution 43: 891–907.
Spence, J.R. (1996). Rosulabryum genus novum (Bryaceae). *The Bryologist * 99: 221–225.
Spence, J.R. (2005). New genera and combinations in Bryaceae (Bryales, Musci) for North America. Phytologia 87: 15–28.
Spence, J.R.; Ramsay, H.P. (2005). New genera and combinations in the Bryaceae (Bryales, Musci) for Australia. *Phytologia * 87: 61–72.
Spence, J.R.; Ramsay, H.P. (2006). Bryaceae, in McCarthy, P.M. (ed.), Flora of Australia. Vol. 51 Mosses 1, pp. 274–348. ABRS, Canberra.
Wang, C.-Y.; Zhao, J.-C. (2009). Phylogeny of Ptychostomum (Bryaceae, Musci) inferred from sequences of nuclear ribosomal DNA internal transcribed spacer (ITS) and chloroplast rps4. Journal of Systematics and Evolution 47: 311–320.
