Schistochilaceae
Corticolous (not in Victoria), on logs, lithophytic or terrestrial, dioicous. Asexual reproduction rarely by leaf fragments. Stems erect to prostrate, irregularly branched, with two ranks of lateral leaves, with or without (not in Victoria) underleaves, sometimes also with laciniate paraphyllia; branches emerging from main stem near an unmodified lateral leaf, with or without (not in Victoria) a collar of tissue at base, or from a lateral leaf composed of only a lobule, without a collar of tissue at base. Lateral leaves incubous to transverse (not in Victoria), contiguous to imbricate, rarely (not in Victoria) shallowly bilobed, concave adaxially, ovate to orbicular, entire and not folded to form a lobule, or almost always with a smaller to larger adaxial lobule, often with one (not in Victoria) or two wings extending from line of attachment between lobe and lobule, and sometimes with additional smaller unistratose ridge-like lamellae; lobe unlobed or rarely pinnately lobed (not in Victoria), ovate, triangular, lanceolate, oblong, elliptic (not in Victoria) or obovate (not in Victoria), entire (not in Victoria) or dentate to ciliate, uni- or multistratose (not in Victoria), with acute, obtuse or rarely piliferous (not in Victoria) or truncate (not in Victoria) apex; lobule ovate, triangular, lanceolate, oblong, elliptic (not in Victoria), rectangular (not in Victoria) or obovate (not in Victoria), entire (not in Victoria) or dentate to ciliate, broadly fused along basiscopic margin with lobe as a keel, uni- or multistratose (not in Victoria), acute, emarginate (not in Victoria) or truncate (not in Victoria), apex free from or united to (not in Victoria) lobe; wings when present uni- or multistratose (not in Victoria), entire, dentate or ciliate; lamellae when present on abaxial surface of lobe and usually also adaxial surface of lobule, entire, dentate or ciliate (not in Victoria). Underleaves when present unlobed (not in Victoria) and elliptic to ovate or orbicular, shallowly to deeply bifid or 4–6-lobed, with entire (not in Victoria), dentate (not in Victoria) or ciliate margins, one to each pair of lateral leaves, smaller than lateral leaves, narrower (not in Victoria) or wider than stem, contiguous to imbricate, sometimes with lamellae adaxially extending beyond sinus at lobe bases toward underleaf base. Leaf cells circular or elliptic to polygonal, smooth or rarely with a domed papilla (not in Victoria), a conical 1–2 cell projection (not in Victoria) or a compound spine multiple cells tall (not in Victoria), thin-walled, rarely differentiated toward margins to form a distinct hyaline border (not in Victoria), without or with small to large trigones, with 2–numerous smooth, granular or granular-botryoidal ovoid to ellipsoid or fusiform oil bodies. Rhizoids usually scattered along stem and branches, occasionally in fascicles from underleaves or lateral leaf lobe bases, without internal peg-like thickenings, purple or hyaline (not in Victoria) to brown (not in Victoria). Androecia terminal on main stem, with bracts and bracteoles similar to vegetative leaves, both bearing 1–numerous antheridia. Sporophyte terminating main stem or short lateral branches, surrounded by shoot calyptra developing into a clavate to cylindric coelocaule, sometimes also surrounded by a perianth; perianth obloid or urceolate to campanulate, with longitudinal lamellae, laciniate at apex. Capsules ellipsoid to cylindric, uni- (not in Victoria) or 3–5-stratose, dehiscing by 4 valves, barely emergent to elevated well above perianth; elaters present, uni- (not in Victoria), bi- or trispiral (not in Victoria). Spores globose, granulate to vermiculate, papillose or echinate (not in Victoria).
One genus and 88 species shared between southern South America, Juan Fernandez, tropical and southern Africa, Madagascar, Sri Lanka, Malesia north to Japan and east to the tropical Pacific and New Zealand and eastern Australia (Schuster & Engel 1977, 1985; Söderström et al. 2016); two species in Victoria.
Gametophytic complexity reaches its peak in the Schistochilaceae and some have been dubbed the most beautiful of all liverworts (Schuster & Engel 1977, 1985). Variation in the elaborate leaf structure in this family provides a wealth of characters for differentiating species with relative ease and contrasts markedly with most leafy liverworts for which differentiating species and establishing infrafamilial classifications has been complicated by a dearth of characters and reliance on often subtle characters (Schuster & Engel 1977, 1985). Schistochila appendiculata (Hook.) Dumort. Ex Trev. of New Zealand is also the largest of the leafy liverworts, supposedly with shoots that can grow to 2.6 cm wide and 110 cm long (Schuster & Engel 1977, 1985). The Victorian species are also large and apart from their highly distinctive leaf morphology, comprising an adaxial lobule and wings attached to the keel between lobe and lobule, are easily distinguished by their large size alone.
The genera Pachyschistochila, Paraschistochila and Pleurocladopsis were recognised as distinct from Schistochila in Schistochilaceae based in part on hyaline rhizoids, a lack of underleaves, and lateral leaves not folded to form lobules respectively. These genera form lineages derived from Schistochila and have now accordingly been subsumed into Schistochila (He & Glenny 2010; He & Sun 2013). Likewise, the monotypic New Caledonian family Perssoniellaceae that was thought to be the closest related family to Schistochilaceae (Schuster 1964), was also shown to have been derived from Schistochila and is now placed in Schistochila (He & Glenny 2010). The distinctive features that defined Perssoniellaceae, including a combination of distichous shoots and large elaborate trigones, have been suggested to be adaptions to epiphytic habitats from terrestrial ancestors (He & Glenny 2010).
He, X. & Glenny, D. (2010). Perssoniella and the genera of Schistochilaceae: a new classification based on molecular phylogenies. Australian Systematic Botany 23: 229–238.
He, X. & Sun, Y. (2013). Multigene evidence reveals the systematic position of Pleurocladopsis simulans (C. Massal.) R.M. Schust. within Schistochila Dumort., Schistochilaceae. Polish Botanical Journal 58: 467–474.
Schuster, R.M. (1964). Studies on Antipodal Hepaticae VI. The suborder Perssoniellinae: morphology, anatomy and possible evolution. Bulletin of the Torrey Botanical Club 91: 479–490.
Schuster, R.M. & Engel, J.J. (1977). Austral Hepaticae V. The Schistochilaceae of South America. Journal of the Hattori Botanical Laboratory 42: 273–423.
Schuster, R.M. & Engel, J.J. (1985). Austral Hepaticae V(2). Temperate and subantarctic Schistochilaceae of Australasia. Journal of the Hattori Botanical Laboratory 58: 255–539.
Söderström, L., Hagborg, A., von Konrat, M., Bartholomew-Began, S., Bell, D., Briscoe, L., Brown, E., Cargill, D.C., Costa, D.P., Crandall-Stotler, B.J., Cooper, E.D., Dauphin, G., Engel, J.J., Feldberg, K., Glenny, D., Gradstein, S.R., He, X., Heinrichs, J., Hentschel, J., Ilkiu-Borges, A.L., Katagiri, T., Konstantinova, N.A., Larraín, J., Long, D.G., Nebel, M., Pócs, T., Puche, F., Reiner-Drehwald, E., Renner, M.A.M., Sass-Gyarmati, A., Schäfer-Verwimp, A., Moragues, J.S., Stotler, R.E., Sukkharak, P., Thiers, B.M., Uribe, J., Váňa, J., Villarreal, J.C., Wigginton, M., Zhang, L. & Zhu, R. (2016). World checklist of hornworts and liverworts. Phytokeys 59: 1–828.
Spinning