Jungermanniaceae Rchb.
Jamesoniellaceae He-Nygrén, Juslén, Ahonen, Glenny & Piippo in He-Nygrén et al., Cladistics 22: 27. 2006.
Type: Jungermannia L.
Plants anisophyllous, usually procumbent or creeping, more rarely erect, opaque, chlorophyllose, and mostly forming red, vinaceous and/or brown wall pigments, medium to large and robust. Branching irregular, sparing, the branches for the most part of Frullania type and lateral-intercalary type (the intercalary branches dorsally displaced in Andrewsianthus, often ventrally displaced); ventral-intercalary branches in a few species; Radula -type basiscopic branches in some taxa; stolons present in some species, geotropic. Stems with cortex weakly differentiated, not forming a hyaloderm. Rhizoids scattered on ventral surface of the stem, rarely more frequent near the ventral leaf base, rarely (Notoscyphoideae) on underleaf bases. Leaves with insertion varying from nearly transverse (but with ventral half usually succubous) to strongly succubous, extending to the stem midline dorsally (beyond in Gottschelia and some Jamesonielloideae; a leaf-free strip present only in Vanaea), with apices undivided or bilobed, less often 3–4-lobed; margins entire (except Chandonanthus) or sometimes armed at bases. Cells usually with trigones small to bulging, at times firm-walled; surface often striate-papillose. Oil-bodies usually coarsely granular to papillose, rarely homogeneous. Underleaves either absent or small, rarely to 0.4× leaves in size (larger in Pseudocephaloziella), lanceolate (bilobed in Notoscyphus). Asexual reproduction, if present, by gemmae.
Dioecious or paroecious, rarely autoecious. Androecia on leading shoots, terminal but becoming intercalary; antheridia 1–3 per bract, the stalk 1–2-seriate. Gynoecia on leading shoots; bracteole to 0.6× bract area, often connate with bracts on one or both sides, or vestigial or absent. Perianth normally present, varying from long and long-emergent (then normally lacking a subtending Isotachis -type perigynium or low marsupium) to abbreviated and ± included within the bracts (then the perianth is normally subtended by a well-developed stem perigynium); perianth usually variously plicate (at least near the mouth), occasionally eplicate, the mouth usually contracted.
Seta usually of numerous cells (reduced in Gerhildiella). Capsule ovoid to short-ellipsoidal, the wall 2–4(7)-stratose; outer layer of cells with nodular thickenings; inner layer of cells with semiannular bands.
Spores papillose-short-vermiculate to verrucate (coarsely and irregularly warty) or baculate. Spore:elater diam. ratio usually ca. 2:1 (ca. 4:1 in Andrewsianthus). Elaters bispiral.
A diverse, polytypic family with eight subfamilies, outlined as follows; note that many of the genera are monotypic.
1) subfam. Chandonanthoideae Inoue has three genera: Chandonanthus Mitt., monotypic, which occurs in temperate Australasia; extra-Austral genera are Plicanthus R.M.Schust. (5 species) and Tetralophozia (R.M.Schust.) Schljakov (3–4 species);
2) subfam. Lophozioideae Macvicar has 12 genera; the Austral genera include:
a) Anastrophyllum (Spruce) Steph., which is widespread in south temperate areas (Engel and Braggins [1998] treat the south temperate species);
b) Andrewsianthus R.M.Schust., with several species in south temperate areas;
c) Gymnocoleopsis (R.M.Schust.) R.M.Schust. has three species, one, G. cylindriformis (Mitt.) R.M.Schust., occurs on Kerguelen Island and Prince Edward Island; a second, G. capensis (S.W.Arnell) R.M.Schust., is from South Africa; a third species, G. multiflorus (Steph.) R.M.Schust., is montane/alpine in the northern Andes and tropical Africa;
d) Lophozia (Dumort.) Dumort. is widespread in south temperate and subantarctic areas;
e) Roivainenia Perss. (cf. Persson and Grolle, 1961) is monotypic and occurs on South Shetland Islands (Bednarek-Ochyra et al., 2000), South Georgia, Falkland Islands (Engel, 1990a) and southern South America (Magellanian and Valdivian regions; see range in Engel, 1990a);
f) Sphenolobopsis R.M.Schust. & N.Kitag., with S. pearsonii (Spruce) R.M.Schust. & N.Kitag., occurs in oceanic western Europe, southern Appalachians, NW North America and on Tristan da Cunha (S. kitagawae R.M.Schust. occurs in Borneo and New Guinea).
Extra-Austral genera are Anastrepta (Lindenb.) Schiffn. (monotypic); Gerhildiella Grolle (monotypic); Gymnocolea (Dumort.) Dumort., with a few species; Hattoria R.M.Schust. (monotypic); Pseudocephaloziella R.M.Schust. (monotypic); and Sphenolobus (Lindb.) Berggr.
3) subfam. Jungermannioideae has 11 genera, four of them Austral:
a) Arctoscyphus Hässel, with two species, both from southern South America (cf. Hässel, 1990; Schuster, 2002a);
b) Cryptostipula R.M.Schust., monotypic, of New Zealand (the status of this genus is uncertain and will be evaluated and discussed in volume 2 of this series);
c) Invisocaulis R.M.Schust., monotypic and known only from Prince Edward Island (Schuster, 2002a);
d) Solenostoma Mitt., with several species in southern temperate areas.
Extra-Austral genera are Cryptocolea R.M.Schust. (monotypic); Cryptocoleopsis Amakawa (monotypic); Diplocolea Amakawa (monotypic); Horikawaella S.Hatt. & Amakawa (monotypic); Jungermannia L. (s. str., with 7–8 species); Nardia Gray; and Phragmatocolea Grolle (monotypic).
4) subfam. Jamesoniellioideae Inoue has eight genera; the Austral ones are:
a) Cryptochila R.M.Schust., with several species in south temperate areas;
b) Cuspidatula Steph. has two species, one in our area;
c) Jamesoniella (Spruce) Carrington has several species in south temperate areas;
d) Nothostrepta R.M.Schust. has two species, N. bifida (Steph.) R.M.Schust. of the Falkland Islands, southern South America and Juan Fernandez, and N. longissima (Steph.) R.M.Schust. of southern South America (Engel, 1978, 1990a; Schuster, 1980c, 2002a);
e) Pisanoa Hässel, monotypic, of southern South America (between 51° and 52° S in wet, coastal southern Chile) (Hässel, 1989; Schuster, 2002a).
Extra-Austral genera are Anomacaulis (R.M.Schust.) R.M.Schust. (monotypic); Denotarisia Grolle (monotypic); and Vanaea (Inoue & Gradst.) Inoue & Gradst. (monotypic). Inoue (1966) included Syzygiella and Syzygiella subg. Protosyzygiella in the Jamesonielloideae. Schuster (1980b, p. 334) elevated the last to generic rank and included both in subfam. Syzygielloideae R.M.Schust., of the Plagiochilaceae. Schuster (2002a, p. 402) included Syzygiella and Protosyzygiella in a key to the genera of the Jamesonielloideae, but stated (p. 402) “although Inoue placed these genera in Jamesonielloideae I think a closer affinity is to the Plagiochilaceae ...” Recent molecular analyses (e.g., Hentschel et al., 2006; He-Nygrén et al., 2006) demonstrated that the affinities of these genera are not with members of Plagiochilaceae, but rather their phylogenetic affinities lie elsewhere. Until further molecular studies are done on these genera, we follow Inoue (1966) in placement of these taxa in subg. Jamesonielloideae. Protosyzygiella (Inoue) R.M.Schust. has one species, P. pseudoconnexa (Herzog) R.M.Schust., of southern South America. Syzygiella has ca. 20 species, and is mostly Neotropical.
5) subfam. Gottschelioideae R.M.Schust. has one genus, Gottschelia Grolle, with two species, and is extra-Austral.
6) subfam. Scaphophyloideae R.M.Schust. has one genus, Scaphophyllum Inoue, monotypic, and is extra-Austral.
7) subfam. Notoscyphoideae R.M.Schust. has one genus, Notoscyphus Mitt., with two species; N. belangerianus (Lehm. & Lindenb.) Mitt. occurs in Cape Prov., South Africa.
8) subfam. Eremonotoideae R.M.Schust. has two genera, both monotypic and extra-Austral, Eremonotus Lindb. & Kaal. ex Pearson and Paramomitrion R.M.Schust.
Rhodoplagiochila R.M.Schust. has only R. rosea R.M.Schust. of Venezuela (Estado Merida, Sierra de Santo Domingo, 3700–3750 m) and was placed by Schuster (1978, 1980b) in Plagiochilaceae. Inoue (1984) placed the genus in “Lophoziaceae,” and Gradstein et al. (2001) followed with a placement in Jungermanniaceae. The genus remains poorly known and subfamilial placement has not been made.
Bragginsella R.M.Schust., a monotypic genus of New Zealand, was placed by Schuster (1997a) in subfam. Jungermannioideae, but, based on the discovery of fertile material, was assigned by Glenny and Malcolm (2005) to Geocalycaceae (see volume 2 of this series for further discussion).
