Racopilum cuspidigerum, photographed by Heino Lepp.

Belongs within: Bryophyta.
Contains: Fontinalaceae, Hypnales, Hookeriales, Rhizogoniaceae, Hypnodendron, Orthotrichaceae, Splachnales, Hedwigiales, Bryaceae, Bartramiaceae, Mniaceae.

The Bryidae are one of the major subgroups of mosses. They are distinguished by the possession of a diplolepideous-alternate peristome of articulated cell wall remnants forming two alternating concentric rings of teeth with cilia often present between the inner teeth (Cox et al. 2000). Bryid mosses have been divided between the pleurocarpus Hypnaneae and the acrocarpous Bryanae, with the Rhizogoniales morphologically intermediate between the two. However, the Rhizogoniales are probably paraphyletic with regard to the Hypnaneae, and Bryanae may be paraphyletic to Rhizogoniales + Hypnaneae combined. Members of the Bryanae include the Bryales, which bear inclined to pendulous, dehiscent capsules on a long seta, with a double peristome with keeled and perforated exostome processes arising from a high basal membrane and alternating with the exostome teeth.

<==Bryidae [Pleurocarpineae]
    |  i. s.: Bryoxiphiaceae [Bryoxiphioideae] O02
    |--Hypnaneae [Pleurocarpae] SK02
    |    |  i. s.: Fontinalaceae D24
    |    |         Leucodontaceae D24
    |    |           |--Habrodon notarisii [incl. H. perpusillus] D24
    |    |           |--Antitrichia curtipendula [=Hypnum curtipendulum] D24
    |    |           |--Pterogonium gracile [=Hypnum gracile; incl. P. ornithopodioides] D24
    |    |           |    |--P. g. var. gracile D24
    |    |           |    |--P. g. var. capense D24
    |    |           |    `--P. g. var. harlecense Jones 1917 D24
    |    |           `--Leucodon Schwägr. 1816 SK02
    |    |                |--L. brachypus D24
    |    |                |--L. canariensis FHH01
    |    |                |--L. immersus D24
    |    |                `--L. sciuroides [=Hypnum sciuroides] D24
    |    |                     |  i. s.: L. s. f. falcata D24
    |    |                     |--L. s. var. sciuroides D24
    |    |                     `--L. s. var. morensis [=Hypnum morense] D24
    |    |--Hypnales SK02
    |    `--Hookeriales SK02
    |--Rhizogoniales [Rhizogonianae] SK02
    |    |--Rhizogoniaceae SK02
    |    |--Hypnodendron SK02
    |    |--Mittenia Lindb. 1863 [Mitteniaceae] SK02
    |    |    `--M. plumula (Mitt.) Lindb. 1863 [=Mniopsis plumula; incl. Mn. rotundifolia, Mittenia rotundifolia] SK02
    |    |--Spiridens Nees 1823 [Spiridentaceae] SK02
    |    |    |--S. aristifolius B57
    |    |    |--S. balfourianus B57
    |    |    |--S. flagellosus B57
    |    |    `--S. vieillardii Schimp. 1865 [incl. S. muelleri] SK02
    |    |--Bescherellia Duby 1873 [Cyrtopodaceae] SK02
    |    |    `--B. elegantissima Duby 1873 (see below for synonymy) SK02
    |    |--Calomnion Hooker & Wilson 1854 [Calomniaceae] SK02
    |    |    |--C. complanatum (Hooker & Wilson) Lindb. 1872 [incl. C. laetum (nom. illeg.)] SK02
    |    |    `--C. milleri Vitt 1991 SK02
    |    |--Pterobryella (Müll.Hal.) Jaeger 1877 [Pterobryellaceae] SK02
    |    |    |--P. breviacuminata Besch. 1878 SK02
    |    |    `--P. praenitens Müll.Hal. 1878 (see below for synonymy) SK02
    |    `--Racopilaceae SK02
    |         |--Powellia Mitt. 1868 SK02
    |         |    `--P. involutifolia Mitt. 1868 [incl. Helicophyllum australe, P. australis (Hampe) Broth. 1907] SK02
    |         `--Racopilum Palisot de Beauvois 1805 SK02
    |              |--R. crinitum Hampe 1870 SK02
    |              |--R. cuspidigerum (Schwägr.) Ångström 1872 SK02
    |              |    |--R. c. var. cuspidigerum SK02
    |              |    `--R. c. var. convolutaceum (Müll.Hal.) Zanten & Dijkstra 1995 (see below for synonymy) SK02
    |              |--R. pacificum Besch. 1898 SK02
    |              |--R. purpurascens Hampe 1876 SK02
    |              |--R. robustum SK02
    |              |--R. spectabile B57
    |              |--R. strumiferum M49
    |              `--R. tomentosum (Hedw.) Brid. 1827 [=Hypopterygium tomentosum] SK02
    `--Bryanae SK02
         |--Orthotrichaceae SK02
         |--Splachnales SK02
         |--Hedwigiales SK02
         `--Bryales SK02
              |--Bryaceae SK02
              |--Bartramiaceae SK02
              |--Mniaceae SK02
              |--Leptostomum Brown 1811 [Leptostomataceae] SK02
              |    |--L. erectum Brown 1811 [incl. Gymnostomum leptostomum] SK02
              |    |--L. inclinans Brown 1811 (see below for synonymy) SK02
              |    |--L. macrocarpon (Hedw.) Bach.Pyl. 1814 [=Bryum macrocarpon] SK02
              |    `--L. menziesii SK02
              |--Aulacomnium Schwägr. 1827 [Aulacomniaceae] SK02
              |    |--A. androgynum [=Mnium androgynum, Orthopyxis androgyna] D24
              |    |--A. palustre (Hedw.) Schwägr. 1827 SK02 (see below for synonymy)
              |    |    |--A. p. var. palustre D24
              |    |    |--A. p. var. fasciculare [=Mnium fasciculare] D24
              |    |    |--A. p. var. imbricatum D24
              |    |    `--A. p. var. laxifolium D24
              |    `--A. turgidum [=Hypnum turgidum, Gymnocybe turgida] D24
              `--Orthodontium Schwägr. 1827 [Orthodontiaceae] SK02
                   |--O. australe Hooker & Wilson 1844 (see below for synonymy) SK02
                   |--O. gracile [=Stableria gracilis] D24
                   |    |--O. g. var. gracile D24
                   |    `--O. g. var. heterocarpa Wats. 1922 D24
                   |--O. inflatum (Mitt.) Paris 1896 (see below for synonymy) SK02
                   |--O. lineare Schwägr. 1827 (see below for synonymy) SK02
                   `--O. pallens (Hooker & Wilson) Broth. 1903 (see below for synonymy) SK02

Nomina nuda: Leptostomum inclinans f. procera Watts & Whitel. 1906 SK02
             Racopilum aeruginosum Müll.Hal. ex Geh. 1877 SK02
             Racopilum convolutaceum f. robusta Broth. & Watts 1915 SK02

Aulacomnium palustre (Hedw.) Schwägr. 1827 SK02 [=Gymnocybe palustris SK02, Mnium palustre D24, Aulacomnium palustre var. eupalustre (nom. illeg.) SK02; incl. A. palustre var. polycephalum D24, Gymnocybe palustris var. ramosa D24]

Bescherellia elegantissima Duby 1873 [incl. Cladomnion bescherelloides (n. n.), Cyrtopus bescherelloides (n. n.), Bescherellia brevifolia Hampe 1876, B. cryphaeoides (Müll.Hal.) Fleischer 1920, B. cyrtopus (n. n.)] SK02

Leptostomum inclinans Brown 1811 [=Gymnostomum inclinans; incl. L. flexipile, L. gracile, L. inclinans var. longiseta Hampe 1856 (n. n.)] SK02

Orthodontium australe Hooker & Wilson 1844 [=Apalodium australe; incl. O. australe ssp. robustiusculum] SK02

Orthodontium inflatum (Mitt.) Paris 1896 [=Apalodium inflatum; incl. Wilsoniella compacta (n. n.), Orthodontium ovale, W. whiteleggei Geh. ex Watts & Whitel. 1906 (n. n.)] SK02

Orthodontium lineare Schwägr. 1827 [=Apalodium lineare (preoc.); incl. O. lanceolatum, A. lanceolatum, A. sulcatum, Orthodontium lineare ssp. sulcatum, O. zetterstedtii] SK02

Orthodontium pallens (Hooker & Wilson) Broth. 1903 [=Weissia pallens, Dicranella pallens, Microdus pallens, Seligeria pallens; incl. Apalodium lineare (nom. illeg.), Zygodon linearis (nom. inv.)] SK02

Pterobryella praenitens Müll.Hal. 1878 [=Dendrohypnum praenitens, Hypnodendron praenitens, Hypnum praenitens] SK02

Racopilum cuspidigerum
var. convolutaceum (Müll.Hal.) Zanten & Dijkstra 1995 [=Hypopterygium convolutaceum, R. convolutaceum (Müll.Hal.) Reichardt 1870, R. convolutum (nom. illeg.); incl. R. australe, R. cristatum, R. laetum, Hypopterygium strumiferum, R. strumiferum] SK02

*Type species of generic name indicated


[B57] Bartram, E. B. 1957. Additional Fijian mosses, III. Journal of the Washington Academy of Sciences 46 (12): 392–396.

Cox, C. J., B. Goffinet, A. E. Newton, A. J. Shaw & T. A. J. Hedderson. 2000. Phylogenetic relationships among the diplolepideous-alternate mosses (Bryidae) inferred from nuclear and chloroplast DNA sequences. Bryologist 103 (2): 224-241.

[D24] Dixon, H. N. 1924. The Student's Handbook of British Mosses 3rd ed. V. V. Sumfield: Eastbourne.

[FHH01] Frey, W., M. Hofmann & H. H. Hilger. 2001. The gametophyte-sporophyte junction: unequivocal hints for two evolutionary lines of archegoniate land plants. Flora 196: 431–445.

[M49] Martin, W. 1949. Distribution of the mosses indigenous to New Zealand (supplement no. 1). Transactions and Proceedings of the Royal Society of New Zealand 77 (3): 355–360.

[O02] Ochyra, R. 2002. Nomenclatural changes in subfamilies of the Dicranaceae. Cryptogamie, Bryologie 23 (4): 345–349.

[SK02] Streimann, H., & N. Klazenga. 2002. Catalogue of Australian Mosses. Flora of Australia Supplementary Series 17. Australian Biological Resources Study: Canberra.

Last updated: 9 January 2018.

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