BFNA Title: Jungermanniaceae
Author: various
Date: Dec. 6, 2017
Edit Level: S
Version: 2 June 29, 2018
Some changes in Mesoptychia

 

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
BFNA Web site: http://www.mobot.org/plantscience/BFNA/bfnamenu.htm

Notice

Return to Home

[For illustrations, see individual genera on Summary page.]

XX. JUNGERMANNIACEAE Rchb.

FNA Editors

Plants forming mats or turfs. Branches  sometimes intercalary from sides of stem, sometimes replacing ventral half of a leaf; usually without flagellae. Leaves alternate, succubous or transverse, plane, simple, emarginate, or 2-lobed, entire; underleaves present or absent, entire or divided for less than half their length. Rhizoids scattered over ventral stem surface. Specialized asexual reproduction of few-celled gemmae, sometimes present on erect gemmiparous branches (absent in most taxa). Gynoecium terminal on an ordinary leafy branch. Perianth with or without subfloral branches, well developed or absent, if present cylindrical, smooth or plicate, mouth very narrow, often abruptly beaked perigynium, but if perianth small or absent then with a deep marsupium.

Genera 7 (5 in the flora), species 34 (18 in the flora): worldwide including Antarctica.

Jungermanniaceae has been much more broadly defined in the past.  It is now clear, however, that morphological similarity between Solenostoma and Jungermannia, and between Mesoptychia and Lophozia, is due to convergence rather than close relationship (Shaw et al. 2015), and a relatively narrowly definition of Jungermanniaceae is necessary.  In view of the close similarity between Solenostoma and Jungermannia, both genera are included in the key below.

SELECTED REFERENCE  Shaw, B., B. Crandall-Stotler, J. Vana, R. E. Stotler, M. von Konrat, J. J. Engel, E. C. Davis, D. G. Long, P. Sova and A. J. Shaw.  2015.  Phylogenetic relationships and morphological evolution in a major clade of leafy liverworts (Phylum Marchantiophyta, Order Jungermanniales): Suborder Jungermanniineae.  Syst. Bot. 40: 27--45. Wagner, D. H. 2013. Rivulariella, gen. nov. (Jungermanniaceae), endemic to western North America. Phytoneuron 2013--10: 1--9.

1.  Stems filiform, leaves transversely inserted, (90--)100--160(--180) /um, lobed for ca. half of their length; oil-bodies, if present, confined to enlarged occeli scattered in the leaf.  1. Eremonotus, p. xxx

1.  Stems not filiform, leaves succubous, 250--3000 /um, unlobed or lobed for no more than 1/3 of their length; oil-bodies present in normal leaf cells.

2.  Leaves 2-lobed (often merely retuse in M. polymorpha, with leaves 250--700 /um and leaf cells lacking trigones). 4. Mesoptychia, p. xxx

2.  Leaves unlobed (often broadly retuse in Liochlaena, with leaves 900--2400 /um and leaf cells with moderately large trigones).

      3.  Leaves ovate to elliptical, cordate, or almost circular, the apex rounded; never  

           producing gemmae.

   4.  Leaves ovate to elliptical or cordate, longer than broad (sometimes almost

           circular and deeply concave in J. polaris); rhizoids confined to stem; perianth

           not beaked.  2. Jungermannia, p. xxx

   4.  Leaves circular to reniform, broader than long; rhizoids sometimes arising from

          leaf bases as well as stem; perianth distinctly beaked.  see Gymnomitriaceae

          (Solenostoma, p. xxx)

     3.  Leaves obovate, rounded or truncate to emarginate or shallowly bilobed; sometimes

          producing erect gemmiparous shoots bearing transverse leaves and large underleaves. 

             5.  Perianth cylindrical, indented at apex; leaf base almost as wide as midleaf width;

                  gemmae 2-celled; plants terrestrial, on wood or organically rich soil. 3.             Liochlaena, p. xxx

             5. Perianth distally plicate, narrowed to apex; leaf narrowed to insertion; gemmae

                  mostly             4-celled; strictly aquatic, attached to rocks.  5. Rivulariella, p. xxx

1. EREMONOTUS Lindberg & Kaalas ex Pearson, Hep. Brit. Isles 200. 1900 * [Greek eremos isolated, deserted, and notos dampness, alluding to the habitat] 

Anomomarsupella R. M. Schuster, Nova Hedwigia 17: 78, pl. 10. 1969

Edwin Urmi

Plants minute. Stems thin, in cross section with only 15--18 firm-walled cells. Lateral leaves transversely inserted, 2-lobed and concave (not plicate as often stated); cells firm-walled, a few of them (irregularly arranged) with a single very large oil body. Underleaves reduced to a slime papilla. Asexual reproduction lacking. Sexual condition dioicous. Androecia wider than vegetative parts. Gynoecia with dorsiventrally flattened perianth. Sporophytes rather massive compared with the gametophytes.

Species 1 (1 in the flora); Arctic North America, Arctic Eurasia.

Selected references: Damsholt, K. 1977: The taxonomic status of Anomomarsupella Schust., (Hepaticae). Lindbergia 4: 132. Hentschel, J., J. A. Paton, H. Schneider, and J. Heinrichs. 2007: Acceptance of Liochlaena Nees and Solenostoma Mitt., the systematic position of Eremonotus Pearsson and notes on Jungermannia L. s.l. (Jungermanniidae) based on chloroplast DNA sequence data. Pl. Syst. Evol. 268: 147--157. Schuster, R. M. 1966--1992: The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian. 6 vol. New York, London, Chicago (Anomomarsupella in vol. 3, pp. 166--170). Urmi, E. 1978: Monographische Studien an Eremonotus myriocarpus (Carring.) Pears. (Hepaticae). Bot. Jahrb. Syst., Pflanzengesch. Pflanzengeog. 99: 498--564.

1. Eremonotus myriocarpus (Carrington) Pearson, Hep. Brit. Isles 201. 1900  F

Jungermannia myriocarpa Carrington, in Carrington and Pearson (ed.), Hep. Brit. Exsicc. (fasc. 2) no. 96. 1879; Anomomarsupella cephalozielloides R.M. Schuster

Plants 2--8 x (0.06--)0.10--0.15(--0.16) mm, sparingly branched, loosely creeping or ascending from a stoloniform system in dense mats, from ocher or dark greenish brown to nearly black (never reddish), somewhat shiny in dry state. Stems 50--75(--80) /um wide, branching lateral intercalary from a single specialized cell in the axils of lateral leaves (Eremonotus type of branching); stems terete, in cross section with 8--10 epidermal and 7--9 central cells; few colourless rhizoids, more and irregularly arranged in the creeping system. Lateral leaves mostly well spaced, appressed or little spreading both wet and dry, concave, in situ (90--)100--160(--180) x (60--)80--120(--130) /um, 2-fid to about the middle; lobes equal, triangular and acute, their margins mostly entire; lobe cells more or less  quadrate in transverse rows, the other less regular and somewhat larger, midleaf cells more or less isodiametric (8--)12--16(--20) /um, surface verruculose, walls evenly thick; oil bodies minute except for a single very large one in some irregularly arranged cells. Ventral leaves vestigial, consisting of 1 cell (a slime papilla). Androecia becoming intercalary, width more than the double of vegetative parts and somewhat dorsiventrally compressed, length and number of bracts indefinite, bracts much larger and more pigmented than vegetative leaves, 2-fid, bulbous and somewhat keeled near the insertion, bracteoles vestigial, 1 single large antheridium, whitish and nearly spherical, with short biseriate stalk, body ca. 120 /um in diameter, filling the whole venter of each bract. Gynoecia on long shoots, sometimes with 1--2 innovations; lateral leaves gradually turning into 3 much larger bracts, free from each other and canaliculate; bracteoles vestigial; perianth longly exerted, (0.4--)0.5--0.6(--0.8) x 0.3 mm, rounded at the apex, with 1 dorsal and 2 ventral folds, mouth shortly 3-lobed and ciliate; 1-layered troughout; with 3--7 archegonia and a few paraphyses. Seta often about 2 mm long and 0.14 mm wide, transverse section with exactly 8 outer and 4 somewhat smaller inner cells. Capsule black and shiny, ellipsoid, dehiscing with 4 valves, wall with 2 cell layers, outer layer with intermediate nodular, inner layer with incomplete half-ring thickenings. Spores brick-red, spheric, (12--)14--16(--18) /um, wall thin, faintly granulated; elaters free, worm-like, ca. 100 x 7 /um, mostly with 2 spirals.

Mostly on humid or wet siliceous rock along streams in sheltered places; often together with Blepharostoma trichophyllum s.l., Solenostoma pumilum s.l., Odontoschisma macounii, Anthelia julacea or Blindia acuta; low to moderate elevations (50--1100 m); Greenland; B.C.; Alaska, Wash.; Arctic Eurasia.

The distribution of Eremonotus myriocarpus in North America is poorly known, due to its nondescript habit. It has often been mixed up with small Marsupella species, but is readily distinguished from these by the lack of red pigments.

2. JUNGERMANNIA Linnaeus, Sp. Pl., 1131. 1753 * [For Ludwig Jungermann, 1572--1653, German botanist]

Vadim Bakalin

Solenostoma (Mitten) Mitten subg. Luridae (Spruce) Müller Frib.

Plants 5--150  x 0.3--4.5 mm,  loosely prostrate to ascending, erect, yellowish or brownish to brown, greenish brown, or deep green, always lacking purple pigmentation. Stem mostly dorsiventrally compressed, cross section elliptic, 100--500 \um wide, 80--480 \um high, branching terminal or intercalary, stolons common in about the half of species; dorsal surface cells subisodiametric, subquadrate, rectangular to shortly elongate, cuticle mostly distinctly striolate; stem cross section composed of uniform cells or of  2 strata the outer of somewhat thicker-walled cells. Rhizoids sparse, yellowish to brownish, never red or purple, decurrent in indistinct fascicles. Leaves inserted ca. 20--60º to axis, sometimes shortly decurrent dorsally, ventrally subtransversely inserted, insertion line mostly arcuate in well-developed plants; blade entire and unlobed; mid leaf cells subisodiametric to slightly oblong, walls thin to slightly or moderately thickened, mostly brownish, trigones distinct and concave or lacking; cuticle finely papillose to striolate at least at stem base. Sexual condition dioicous or paroicous. Perianth terminal on main axis with ca. 5--15 archegonia, on main axis, mostly 2(--3)-stratose in proximal 2/3, cells in mid perianth subisodiametrical to shortly rectangular, never linear, innovations frequent, 1--2 per perianth, perianth emergent for 1/2--2/3\x its length, fusiform to clavate, gradually narrower or abruptly constricted to the mouth, sometimes 4--5-plicate distally, perianth mouth crenulate; perigynium indistinct; bracts slightly larger than sterile leaves, in 1 pair. Androecia intercalary, 2--10 pairs of bracts, with 1(--2) antheridia per bract, stalk of 2 rows of cells, bracts subimbricate, inflated in proximal half and imbricate distally. Capsule wall 2-stratose. Elaters with or without homogenous endings. Spores 12--18 \um, finely papillose, reddish brown, spheric.

Species 9 (6 in the flora): North America, Central America, South America, Eurasia, Africa, Pacific Islands, Antarctica.

The genus Jungermannia was established by Linnaeus (1753) and for long included all species of leafy liverworts. Over time the concept has become much narrower. Adopted here is R. N. Schljakov’s (1981) narrow generic concept, which matches that of R. M. Schuster’s (1969) Solenostoma subg. Luridae.

SELECTED REFERENCES  Damsholt, K. and J. Váňa. 1977. The genus Jungermannia L. emend. Dumort. (Hepaticae) in Greenland. Lindbergia. 4: 1--26.   Doyle, W. T., and R. E. Stotler. 2006. Contribution toward a bryoflora of California. III. Keys and Annotated Species Catalogue for Liverworts and Hornworts. Madrono.53: 89--197.  Hicks, M. L. 1992. Guide to the Liverworts of North Carolina. Durham, N. C. Schuster, R. M. 1969. The Hepaticae and Anthocerotae of North America. Vol. 2. New York and London.  Schuster, R. M. 1988. The hepaticae of South Greenland. Beih. Nova Hedwigia 92.  Schuster, R. M. and K. Damsholt. 1974. The hepaticae of West Greenland.  Meddel. Grønland 199: 5--373. Schljakov, R. N. 1981. [No title.] Pečen. Mhi Severa SSSR 4: 1--224.   Váňa, J. and W. S. Hong. 1999. The genus Jungermannia in western North America. Lindbergia 24:133--144.

1. Leaves semi-imbricate, loosely sheathing stem at base, plants semi-erect, dioicous, mostly more than 1.5 mm wide, brownish green, deep green to blackish.

2. Plants larger, mostly more 2 mm wide, leaves lax, frequently undulate at margins, distinctly cordate and widely triangular, mid leaf cell walls colorless to brownish … 3. Jungermannia eucordifolia

2. Plants smaller, mostly to 2 mm wide, leaves rigid, not or rarely and obscurely undulate at margins, mostly ovate, mid leaf cell walls orange-brown to rarely brownish … 4. Jungermannia exsertifolia

1. Leaves (except area near perianth and androecia) horizontally spreading (sometimes subimbricate in small prostrate plants that are narrower than 1.2 mm); plants prostrate to ascending, rarely erect in perianthous shoots; paroicous or dioicous, less than 1.5 mm wide, or if larger mostly yellowish brownish to yellowish green in color with distinctly horizontally spreading leaves.

3. Plants paroicous or rarely (in Arctic stations only) heteroicous, freely fertile.

4. Width of stem ca. 1/3(--1/4)\x plant width, plants commonly black to blackish; mostly Arctic … 5. Jungermannia polaris

4. Width of stem less than (1/4--)1/5\x plant width; plants mostly green to brownish green; widespread … 6. Jungermannia pumila

3. Plants dioicous, rarely fertile.

5. Plants yellowish to greenish brown, rarely deep green, mostly more than 1.5 mm wide; leaves distinctly horizontally spreading; boreal to temperate areas, rarely in tundra … 1. Jungermannia atrovirens

5. Plants blackish to green-black, greenish brown, less than 1 mm; leaves subimbricate; mostly Arctic or alpine… 2. Jungermannia borealis

1. Jungermannia atrovirens Dumortier, Syll. Jungerm. Europ., 51. 1831

Solenostoma atrovirens (Dumortier) Muller Frib.; Jungermannia tristis Nees

Plants 12--20 x 1.4--2 mm, loosely prostrate to ascending, yellowish, brownish to brown and greenish brown, rarely in shade) deep green. Stem 175--315 \um wide, 120--210 \um high, branching rare, terminal or intercalary; dorsal surface cells 60--125 x 22--30 \um, mostly oblong-rectangular, cuticle distinctly striolate; stem cross section outer cells larger than the inner, ca. 20--23 \um, walls thickened, brownish to brown, inner cells ca. 12--15 \um, mostly pentagonal, less thick-walled, with walls yellowish to colorless, trigones small and concave. Rhizoids sparse, yellowish, decurrent on the stem. Leaves inserted at ca. 40º to axis, decurrent dorsally, ventrally subtransversely inserted, insertion line arcuate, shortly decurrent; 1125--1425 x 825--1125 \um, 0.8--0.9:1, ovate to ovate-triangular, flattened, rarely slightly concave-channeled; mid leaf cells 25--35 x 22--30 \um, subisodiametric to slightly oblong, walls mostly thickened, brownish or yellowish, trigones distinct concave; cuticle papillose throughout. Oil-bodies 2--3(--4) per cell, spherical, 4--7 \um to fusiform and ellipsoidal, 7--10(--15) x 4--7 \um. Sexual condition dioicous. Perianth terminal with ca. 10 archegonia, on main axis, innovations frequent, about 1 per perianth, originating below bracts; perianth emergent for 1/2--2/3\x length, ca. 1700 x 700 \um, fusiform to clavate, mouth gradually narrower or abruptly constricted to the beaked mouth, sometimes 4--5-plicate distally, perianth mouth crenulate; perigynium indistinct; bracts slightly larger than sterile leaves, in 1 pair. Androecia intercalary, in 8--10 pairs of bracts, with 1 antheridium per bract, bracts subimbricate, inflated in proximal half and imbricate distally. Elaters ca. 8 \um wide. Spores 14--18 \um in diameter, finely papillose, reddish brown.

Moist soil and rocks near streams, mostly in forested areas; 0--2800 m; Greenland; Alta., B.C., N.S., Que., Alaska, Calif., Colo., Idaho, Mont., Oreg., Utah, Wash.; Eurasia; Africa

2. Jungermannia borealis Damsholt & Váňa, Lindbergia 4: 5. 1977

Plants 7--15 x 0.4--0.55 mm, perianthous plants to 1 mm wide, ascending, blackish green or black-brown, rarely brown and greenish brown, becoming green apically. Stem ca. 100--150 \um wide, 120--200 \um high; dorsal surface cells ca. 30--55 x 12--22 \um, walls brownish; stem cross section outer cells ca. 22--27 \um, slightly thickened, walls brownish, inner cells ca. 12--18 \um. Rhizoids sparse, colorless, grayish to gray-brown, in indistinct fascicles, mostly 90º to stem. Leaves distant, contiguous to subimbricate, inserted ca. 80--90º to stem, 400--550 x 350--550 \um, 0.9--1:1, widely ovate to subtriangular, concave to concave-channeled; mid leaf cells ca. 17--25 x 15--20 \um, mostly hexagonal, walls thin, brownish, trigones triangular to convex, marginal cells 10--15 \um, with more deeply colored and thickened walls; cuticle smooth throughout. Oil bodies 2--9(--12) per cell, sub spherical, ca. 408 \um in diameter, to ellipsoidal or irregular oblong, 6--14 x 4--8 \um, finely granulate. Sexual condition dioicous. Perianth terminal with ca. 10 archegonia within, on main axis, innovations frequent, 1 per perianth, originating below bracts; perianth emergent 1/2--2/3\x its length, 1750--1925 x 630--700 \um, fusiform to clavate, gradually narrower or suddenly constricted to beaked mouth, perianth mouth crenulate, distal perianth 10--12 \um, subquadrate to isodiametric, walls brownish, thickened, trigones triangular to convex, medially 25--30 x 15--20 \um, rectangular to obliquely rectangular, proximally 30--50 x 15--23 \um, rectangular, walls thin, colorless, cuticle striolate-papillose except for distal 1/4; perigynium absent; bracts slightly larger than sterile leaves, in 1 pair, ca. 900--1020 x 880--980 \um, ovate to triangular, concave. Androecia intercalary, of 8--12 pairs of bracts, with 1 antheridium per bract, bracts subimbricate, inflated in proximal half and imbricate distally. Capsule wall 2--stratose. Elaters ca. 7--8 \um wide. Spores 14--16 \um, finely papillose, reddish brown.

Shaded rocks and soil near streams, snow bed habitats, mostly unforested areas;  elevation unknown; Greenland; B.C., Nunavut; Alaska, Calif.; Eurasia.

Jungermannia borealis has been poorly understood due to poor type specimen plants and unclear protologues. K. Damsholt and J. Váňa (1977) found all previous treatments in discrepancy with types, and described Jungermannia borealis to provide a legitimate name to avoid misinterpretation of related species.

3. Jungermannia eucordifolia Schljakov, Pečen. Mhi Severa SSSR 4: 37. 1981

Jungermannia exsertifolia Stephani ssp. cordifolia (Dumort.) Váňa; J. exsertifolia subsp. cordifolia var. pendletonii (Pears.) Váňa; Solenostoma cordifolium (Hooker) Stephani

Plants 25--35 x 1.4--3.5 mm, semi-erect, blackish green, deep green-brown, to black. Stem ca. 500 \um wide and ca. 450 \um high, rarely branching; dorsal surface cells 125--200 x 20--30 \um, linear to long-rectangular; stem cross section outer cells undifferentiated, external walls brown, inner cells almost colorless cells, walls thin, sometimes lightly flexuous, trigones small and concave. Rhizoids sparse, brownish, decurrent on stem or erect. Leaves contiguous to distant, 1575--1750 x 1365--1575 \um, 0.8--0.9:1, triangular, widest in proximal 1/3, concave-channeled, clasping the stem; mid leaf cells 38--50 x 22--30 \um, rectangular to oblong-hexagonal, walls thin to somewhat thickened, brownish to brown, trigones absent or very small and indistinct; marginal cells 22--33 x 22--28 \um, isodiametric, trigones distinct, concave; cuticle slightly striolate in proximal 1/4. Oil bodies 2--4(--8) per cell, ellipsoidal to oblong, 7--11(--15) x 4--7 \um, or spherical, 4--7 \um, finely papillose. Sexual condition dioicous. Androecia intercalary, spicate, with ca. 10 pairs of bracts, one generation separated from another by 1--2 pairs of sterile leaves, bracts triangular, inflated in the base, with 1 antheridium per bract, ca. 30--35 x 23 \um. Perianth with innovations common, 2--3 per perianth, ventral or lateral, originating in axils of both bracts and sterile leaves; perianth clavate, constricted to the mouth, pluriplicate near mouth, emergent for 1/2\x length, mouth crenulate, perianth 1-stratose in distal 1/3, 2-stratose medially, 3-stratose near base; cuticle striolate, except distal 1/4, cells at mid perianth 38--45 x 15--22 \um, walls thickened, trigones distinct, concave; perigynium absent; bracts just below perianth in 1 pair, slightly larger than sterile leaves, subtransversely inserted, suborbicular, ca. 2625--2730 \um. Capsule ellipsoidal. Elaters 100--140 x 8 \um. Spores 19--24 \um, finely papillose.

Wet rocks near streams, waterfalls, beds of sluggishly flowing streams, ponds, oligotrophic bogs with open places; 0--4000; Greenland; Alta., B.C., Nfld. and Labr., N.S., Ont., Que., Yukon; Alaska, Ariz., Calif., Colo., Conn., Idaho, Maine, Mass., Mich., Mont., Nev., N.H., N.Mex., N.Y., Oreg., Utah, Vt., Wash., Wis., Wyo.; Eurasia.

4. Jungermannia exsertifolia Stephani, Sp. Hepat. 6: 86. 1917  F

Jungermannia cordifolia Hooker ssp. exsertifolia Amakawa,

Plants (3--)15--50(--100) x (0.7--)1--2(--2.5) mm, ascending to erect in dense patches, blackish to brown and greenish brown, sometimes with violet rim along leaf margin, sometimes in herbarium plants becoming reddish brown, branching ventral, mostly as leafless geotropic stolons. Stem 250--350 x 150--250 \um, elliptic in cross section, dorsal surface cells slightly thick-walled, rectangular to oblong-hexagonal, with small concave or indistinct trigones, 60--100 x 23--30 \um, cuticle partly indistinctly striolate; stem cross section with large-celled outer layer with cell walls thickened, brown, ca. 20--26 \um, inner layer of smaller, more or less thin-walled, rectangular to oblong-hexagonal cells with small concave to indistinct trigones, ca. 15--25 \um. Rhizoids isolated to sparse, colorless to brownish, in indistinct fascicles or running down the stem. Leaves contiguous to subimbricate, inserted at 40--85° to axis, dorsally subtransverse, not decurrent, ventrally subtransversely inserted, insertion line straight to loosely arched, not decurrent, 600--1750 x 480--1630 \um, 0.85--0.95:1, triangular to triangular-ovate, or ovate, concave, widest at 1/2--1/3\x leaf length in proximal half of the leaf; mid leaf cells thin-walled or slightly but evidently thickened, orange to brown, subisodiametric with lumen rounded to elliptic, 25--30(--38) x 20--25 \um, trigones indistinct to rarely convex; marginal cells 12.5--17.5 \um, with thin to slightly thickened (especially the external) walls, walls more deeply orange-brown, than medial leaf cells; cuticle distinctly papillose in the proximal half of the leaf, in distal portion very indistinctly striolate, or only obscurely striolate in proximal 1/6. Oil bodies granulate, 2--3 per cell, ovate, ca. 8--12 x 7--8 \um. Sexual condition dioicous. Androecia intercalary, with 4--5 pairs of bracts, 1--2 antheridia per bract, bracts triangular, similar to sterile leaves in size, inflated in proximal half, imbricate distally, reddish in inflated area.

Acid and basic rock, along streams, mostly in higher mountain elevations of forest belts; 1200--2200 m; Calif., Mont.; e Asia.

5. Jungermannia polaris Lindberg, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 23: 560. 1867

Solenostoma polare (Lindberg) R. M. Schuster

Plants 1--5 x 0.3--0.7(--0.9) mm, prostrate to ascending (in fertile plants), deep green or brownish to brown-green in deeply colored shoots; fertile plants larger than the sterile. Stem flexuous 175--250 \um in diameter, dorsal surface cells more or less thin-walled, mostly rectangular to obliquely rectangular, 25--50 x 12.5--20 \um, cuticle finely striolate; stem cross section cells more or less uniform, slightly smaller within, more or less thin-walled or indistinctly thickened, with small concave trigones, outer cells ca. 17--23 \um, inner ca. 15--20 \um. Rhizoids dense, brownish, in indistinct fascicles or bundles. Leaves distant to contiguous, inserted at 45--75° to axis, dorsally not or barely decurrent, ventrally subtransversely inserted, not decurrent; 300--500 x 350--600 \um, 1:0.7--0.95, reniform to widely triangular, distinctly concave, widest at proximal 1/3\x leaf length; mid leaf cells with slightly thickened walls, brownish, subisodiametric to quadrate, 17--25 x 12--23 \um, trigones small, concave; margin cells 12--20 \um, with brown, slightly thickened walls; cuticle smooth throughout or loosely striolate. Oil bodies (1--)2--4(--5) per cell. Sexual condition paroicous. Perianth brown at narrowest part, emergent for 1/3--4/5\x length, fusiform to obpyriform and oblong-ovate, gradually narrowed to the mouth, surface smooth, lightly pluriplicate at mouth, ca. (1000--)1610 x (500--)700 \um, mouth crenulate, perianth in distally 1-stratose, cells quadrate to rectangular, 15--33 x 11--15 \um, walls thickened, brown, cuticle smooth, at mid perianth, 1--2-stratose, cells oblong to obliquely rectangular, 25--40 x 12--17 \um, walls thickened in colored area and thin in colorless parts, cuticle striolate in proximal portion, perianth in proximal portion 2-stratose, cells rectangular to oblong-elliptic, 45--53 x 15--18 \um, walls thin; perigynium indistinct or very low, ca. 8--10 archegonia in perichaetium; bracts just below perianth in one pair, similar to sterile leaves, but frequently undulate marginally, elliptic to orbicular, widest in the mid portion, 0.63--0.77 x 0.7--0.735. Androecia below perianth, separated from the female bracts by the one pair of sterile leaves, with 1(--2) pair of bracts, with 1(--2) antheridium per bract, ca. 65 x 25 \um; bracts strongly inflated basally, partly violet in inflated area, apex deflexed, ca. 420 x 480--600 \um. Capsule dark brown, subspherical to shortly ovoid, outer cells subrectangular, 12014 \um in width, with 2--4 nodular thickenings, 0--2 on horizontal wall; inner layer narrowly rectangular, with 4--6 annular bands. Elaters ca. 7--8 \um wide. Spores 15--18 \um, finely papillose, yellowish brown.

Moist rocks and sandstone near streams, fine-grained soil in snow-bed habitats, frequently associated with cryophilous and basiphilous plants like Anthelia juratzkana, Athalamia hyalina and Preissia quadrata; 0--3500 m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), Nunavut, Que.; Alaska, Calif., Mich., Minn., Mont., Wash., Wis.; Eurasia.

6. Jungermannia pumila Withering, Arr. Brit. Pl. (ed. 3) 3: 883. 1796

Jungermannia karl-muelleri Grolle; J. karl-mulleri Grolle ssp. heteroicum (R. M. Schuster) Stotler & Crandall-Stotler; Solenostoma oblongifolium Müller Frib. subsp. heteroicum R. M. Schuster & Damsholt; S. pumilum (Withering) Grolle subsp. anomalum R. M. Schuster & Damsholt; S. pumilum (Withering) Müller Frib.

Plants 3--5 x 0.81--1.2(--2) mm, ascending to erect, dirty green, yellowish green to deep green. Stem 140--210 \um wide, 100--145 \um high; dorsal surface cells 20--33 x 25--30 \um, subquadrate to shortly rectangular, cuticle striolate-papillose; cells of stem cross section dorsally larger, ca. 22--30 \um, thick-walled, yellowish, near ventral side smaller both externally and within, walls slightly thickened, brownish. Rhizoids sparse in indistinct fascicles, colorless to grayish and brownish. Leaves distant to contiguous and subimbricate, inserted at 30--35° to axis, but near perianthous plant apex leaves subtransversely inserted, dorsally subtransversely inserted, not or barely decurrent, ventrally inserted subtransversely, insertion line arched, not decurrent; 400--900 x 250--700 \um, 0.7--0.9(--1):1, ovate to rarely sublingulate and ovate-triangle, distinctly concave-channeled, clinging to the stem at base; mid leaf cells moderately thick-walled to thin-walled, 25--38 x 17--24 \um, walls yellowish, trigones small, but distinct, concave; along margin 10--20 \um, slightly thickened, walls yellowish to brownish, cuticle smooth throughout. Oil bodies 2--6 per cell, 4--8(--10) x (17--)20--25 \um, finely granulate. Sexual condition paroicous or very rarely heteroicous, in Arctic stations freely fertile. Perianth fusiform, gradually narrowed to pluriplicate mouth, 910--1400 x 490--630 \um, with 8--10 archegonia per gynoecium, emergent from bracts for 1/2--3/4\x perianth length, distal cells ca. 17--20 \um, isodiametric, 5--6-gonal, trigones distinct concave, cuticle smooth, in middle 1-stratose, cells oblong, ca. 17--25 x 12--15 \um, thin-walled, trigones triangular to concave, cuticle striolate, proximally 2-stratose, 33--50 x 15--22 \um, rectangular to oblique-rectangular and linear, walls thin to thickened, trigones concave, cuticle distinctly papillose; perigynium indistinct; bracts just below perianth in 1 pair, ca. 1190 x 930 \um, similar to sterile leaves. Androecia below perianth in 2(--3) pairs of bracts, ca. 1050 x 930 \um, ovate-triangular, slightly larger than sterile leaves, strongly inflated near the base, with 1 antheridium per bract. Seta ca. 4500 \um. Capsule elliptic, 650--700 x 250--300 \um, 2-layered, inner cells 50--70 x 9--12 \um, with annular thickenings, outer cells 60--70 x 28--30 \um with 6--8 nodular thickenings in vertical wall. Elaters ca. 140--160 x 7--11 \um, with homogenous tapering ends or without such.

Shaded moist soil and rocks near streams mostly in acidic to neutral environments; 2100--2600 m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., N.S., Nunavut, Ont., Que.; Calif., Colo., Conn., D.C., Idaho, Ill., Iowa, Ky., Maine, Md., Mass., Mich., Minn., Mont., N.H., N.J., N.Mex., N.Y., N.C., Oreg., Pa., R.I., Tenn., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Eurasia.

The synonyms of Jungermannia pumila given above are based not on the literature (for example in R. M. Schuster and K. Damsholt 1974), but directly on study of the type specimens. For the description of this taxonomic confusion see K. Damsholt and J. Vana 1977.

3. LIOCHLAENA Nees, Syn. Hepat., 150 1844 [Greek, leios, smooth, and chlaina, cover, alluding to perianth not folded distally]

Vadim Bakalin

Plants 10--40 mm x 2--3(--5) mm, mostly prostrate, rarely ascending (perianthous plants), pale green, yellowish green to deep green, blackish in very wet places, rarely tinted orange, loosely adhering to the substrate by tawny rhizoids. Branching intercalary. Rhizoids often dense, colorless to yellowish or brownish, never red or purplish. Leaves contiguous, inserted at 30--40° with stem, but near apex of perianthous plants leaves subtransversely inserted, dorsally shortly but distinctly decurrent, ventrally inserted subtransversely, insertion line arched, not decurrent; leaves 900--1600(--2400) x 700-1400(-1600) /um, obliquely lingulate to oval, frequently retuse in larger leaves, sheathing the stem proximally, erect-flattened distally; midleaf cells with walls thin to slightly thickened, walls yellowish to colorless, trigones moderate in size, mostly concave, rarely slightly convex; cuticle verrucose, sometimes indistinctly so. Specialized asexual reproduction by gemmae, very rare, ellipsoidal to shortly fusiform, 2-celled.  Sexual condition paroicous or dioicous. Androecia below perianth or in separate branches, of 1--13 pairs of bracts, oblique-lingulate, slightly larger than sterile leaves, strongly inflated near the base, antheridia 1--3 per bract. Perianth terminal on main axis, cylindric to clavate, abruptly constricted to the beaked mouth, apex often concave with beak projecting from the concavity, surface smooth, emergent from bracts for 1/2--3/4 times the perianth length. Capsule oblong-ellipsoidal, wall 2-layered. Elaters 2-spiral, ca. 8--10 \um in width. Spores 12--14 \um, verrucose.

Species 4 (2 in the flora); North America, Eurasia, Pacific Islands.

Liochlaena includes two closely related species sometimes regarded as subspecies of one species. The main distinctions are in inflorescence type, gemmae production and (to a lesser degree) in mid leaf cell size. Liochlaena subulata is distributed mostly in Asia, but area of L. lanceolata mostly confined to Europe and North America.

SELECTED REFERENCES Grolle, R. 1966. Die Typisierung von Jungermannia L. und Jungermannia lanceolata L. Taxon 15: 186--191.  Váňa, J. 1973. Studien über die Jungermannioideae (Hepaticae). 3. Jungermannia subg.Liochlaena. Folia Geobot. Phytotax.. 8: 397--416. Váňa J. and W. S. Hong. 1999. The genus Jungermannia in western North America. Lindbergia 24: 133--144.

1. Plants paroicous, freely fertile, gemmae virtually absent or very rare, midleaf cells 35--65 x 30--45 \um  … 1. Liochlaena lanceolata

1. Plants dioicous, rarely fertile, gemmae common, midleaf cells 25--40 x 25--35\um … 2. Liochlaena subulata

1. Liochlaena lanceolata Nees, Nat. Pflanzenfam., 150. 1845  F

Jungermannia leiantha Grolle

Midleaf cells 35--65 x 30--45 \um . Oil bodies (4--)6--11(--14) per cell, spheric or oblong, 5--13 x 5--7 \um. Gemmae very rare, ellipsoidal. Sexual condition paroicous; frequently fertile. Androecia below perianth, with 1--5 pairs of bracts.

Shaded soil, decaying wood, forests, mostly coniferous; 0--3500 m; Alta., B.C., Nfld. and Labr., Ont., Que., Ala.; Alaska, Ariz., Calif., Colo., Conn., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine, Mass., Mich., Minn., Mont., Nev., N.H., N.J., N.Y., N.C., Oreg., R.I., Tenn., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Eurasia.

Liochlaena lanceolata has long been classified under the name Jungermannia lanceolata L.  According to Grolle (1966), the type of this name is a different species (i.e. Jungermannia atrovirens Dumort.), and it is not correct to apply it to Liochlaena lanceolata. The chromosome number is 16--18.

2. Liochlaena subulata (A. Evans) Schljakov, Pečen. Mkhi severa SSSR, 71. 1981

Jungermannia subulata A. Evans, Trans. Connect. Acad. Arts Sci. 8: 258. 1892

Jungermannia lanceolata ssp. stephanii Amakawa Journ. Hattori Bot. Lab. 22: 71. 1960

Midleaf cells 25--40 x 25--35\um. Oil-bodies 6--8 per cell, spheric or oblong, 5--17 x 5--7 \um. Gemmae ellipsoidal to shortly fusiform colorless to brownish, commonly present on specialized attenuate shoots with scale-like, eroded leaves. Sexual condition dioicous. Rarely fertilized. Androecia in separate branches, semispicate, intercalary, with 3--12 pairs of bracts.

Moist decaying wood, wet soil; ~500-1500; Minn., Miss.; Eurasia, Pacific Islands.

The chromosome number of Liochlaena subulata is 9.

4. MESOPTYCHIA (Lindberg) A. Evans, Ottawa Naturalist 17: 15 1903 [Greek mesos, middle, and ptychos, plica, alluding to weakly plicate leaves]

Vadim Bakalin

Jungermannia sect. Mesoptychia Lindberg, Kongl. Svenska. Vetensk. Acad. Handl., n.s. 23 (5): 39. 1889; Leiocolea (Müller Friburg) H. Buch; Lophozia subgen. Leiocolea Müller Friburg 

Plants 3--90 x 0.4--5 mm, prostrate to loosely ascending or rarely erect in dense patches, brown to green-brown or yellowish brown. Stem oval in section, 50--500 \um in width and 40--470 \um in height, outer cells thick-walled, inner cells thinner-walled, or (in Leiocolea badensis) composed of uniform cells. Rhizoids brownish to colorless, at right angles to stem, in tufts or tawny fascicles or scattered. Leaves contiguous to distant, subimbricate in some Arctic phases, very obliquely to almost horizontally inserted, dorsally not or short-decurrent, ventrally subtransversely inserted, 2-lobed for 1/8--1/3 of leaf length to entire and emarginate, sinus angular, crescentic or gibbous, lobes arcuate to biconcave, acute or rarely rounded. Cells in midleaf mostly elongate, walls thin, trigones large and convex to small, concave-sided, and indistinct, cuticle coarsely papillose to almost smooth. Oil-bodies brownish, granulate, without central eye, 2--12 per cell. Underleaves  rarely obsolete, commonly lanceolate and lacinulate, undivided or 2- or 3-fid, frequently with acute teeth or (more rarely) ciliate; rarely reduced to slime papillae. Specialized asexual reproduction absent or by ellipsoidal gemmae. Sexual condition dioicous or monoicous. Androecia with bracts not strongly inflated at base, with an additional tooth near dorsal margin; antheridia 1--2(--3) per bract. Perianth terminal on main stem, growth of stem rarely continuing by adventitious branches proximal to perianth; bracts proximal to perianth in 1 pair; perianth tubular to tubular-subclavate, smooth or shallowly plicate, mostly abruptly tightened to the beaked mouth, emergent from bracts for 1/2--3/4 of length, mouth crenulate to dentate, teeth 1--2(--3)-celled, of elongate cells; perigynium absent or (rarely) present. Capsule wall (2--)3--4-stratose. Elaters 2-spiral.

Species 16 (8 in the flora): North America, Eurasia.

SELECTED REFERENCES Hong W.S. 2002. Leiocolea in Western North America. Lindbergia 27: 97--103. Schuster R.M. 1988. The Hepaticae of South Greenland. Beih. Nova Hedwigia. Heft 92. 255 pp. Schuster R.M. 1969. The Hepaticae and Anthocerotae of North America East of the Hundredth Meridian, Vol. 2. Columbia University Press, New York.

1. Plants 3--6 mm wide; leaves widest in proximal third.

2. Plants paroicous, secondary pigmentation brown, underleaves divided into 2--3 laciniae bearing many cilia at narrow angle to lacinia margins, perigynium absent … 7. Mesoptychia rutheana

2. Plants dioicous, secondary pigmentation purple, underleaves mostly 2-lobed into two laciniae, with margins bearing numerous cilia spreading at right angle with margin, perigynium well developed, strongly rhizogenous, at right angle to stem … 8. Mesoptychia sahlbergii

1. Plants 0.3-2.8 mm wide; underleaves obsolete, laciniate, or 2-lobed to near base, cilia few or absent, mostly with short teeth confined to underleaf base, or entire; leaves widest near middle, rarely in proximal third.

3. Leaves with smooth or very obscurely papillose cuticle, plants not translucent, wider than 1 mm, underleaves common, midleaf cells isodiametrical or nearly so, commonly with strongly convex trigones, ellipsoidal gemmae frequently present … 5. Mesoptychia heterocolpos

3. Leaves with moderately to coarsely papillose, rarely obscurely papillose cuticle (rarely almost smooth, but in the latter case plants translucent, narrower 1 mm wide, underleaves obsolete or 1--5 cells long), midleaf cells mostly with concave trigones, oblong; gemmae absent.

4. Plants translucent, 0.3--0.6 mm wide, underleaves obsolete to small, up to 5 cells long, mostly in the form of 1-seriate cilia, freely deciduous; oil bodies mostly spherical, finely papillose, colorless to grayish, small, not filling cell lumen; perianth beaked or not; capsule wall 2--3-stratose (where known) … 5

5. Plants paroicous, although archegonia never observed fertilized, leaves mostly entire to emarginate, rarely bilobed … 6. Mesoptychia polymorpha

5. Plants dioicous, archegonia freely fertilized, leaves clearly bilobed, with acite or nearly so lobes 1. Mesoptychia badensis

4. Plants not translucent, 1--2.8 mm wide, underleaves regularly present, but frequently hidden in tawny rhizoids, not deciduous, 2- to variously lobed at the base; oil bodies in leaf cells mostly oblong, brownish, finely granulate, commonly filling cell lumen; perianth beaked; capsule wall 3--4-stratose.

6. Plants monoicous, calciphilous or neutral habitats … 4. Mesoptychia gillmanii

6. Plants dioicous, strictly calciphilous.

7. Midleaf cells 29--45 \um wide, leaves asymmetrical; plants 1.8--2.8 mm wide … 2. Mesoptychia bantriensis

7. Midleaf cells 22--35 \um wide; leaves almost symmetrical; plants 1.6--1.9 mm wide … 3. Mesoptychia collaris

1. Mesoptychia badensis (Gottsche) L. Söderström & Váňa, Phytotaxa 65: 52. 2012

Jungermannia badensis Gottsche, Hepat. Eur., 95. 1859; Lophozia badensis (Gottsche) Schiffner; L. badensis var. apiculata R.M. Schuster; Leiocolea badensis (Gottsche) Jørgensen

Plants 3--8 x 0.3--0.6 mm, prostrate to ascending and semi-erect when growing between mosses, brownish green with more deeply colored apical portion of perianth and female bracts.  Stem 100--180 \um in diameter, in section cells more or less uniform, with thin or moderately thick walls, 15--22 \um in diameter, external wall of stem thicker.  Rhizoids colorless, in tufts or scattered.  Leaves contiguous to distant, almost symmetrical, rectangular, inserted at ca. 30--50° with axis, dorsally not decurrent, ventrally subtransversely inserted, 270--450 x 250--420 \um, 0.9--1.1 times as long as wide, plane, lobed 1/4--1/3 of length, sinus rectangular, lobes triangular, blunt.  Cells in midleaf 25--45 x 25--30 \um, walls thin, trigones medium in size, triangular to convex; marginal cells 20--30 \um, thin-walled;   cuticle distinctly papillose near ventral leaf base, almost smooth in distal half of leaf.  Oil-bodies 2--5(--7) per cell, spherical and 5--8 \um to (rarely) ellipsoidal, to 14 \um.  Underleaves irregularly present and easily deciduous, mostly 2--3 cells in length, ca. 40--45 \um.  Specialized asexual reproduction absent.  Sexual condition dioicous.  Androecia intercalary on small shoots, bracts in 3--4 pairs,  concave to cupped and channeled, commonly with one tooth near dorsal leaf base, antheridia 1 per bract, bracteole absent or the same size as the stem underleaves.  Perianth tubular to tubular-clavate, smooth to loosely folded, suddenly tightened to the beaked mouth,  emergent from bracts for 1/2--2/3 of its length, mouth crenulate to dentate,  frequently lacerate;   trapezoidal,  2-lobed for 1/6--1/5 of the length, sinus gibbous. 

Obligate calciphile, growing on calcium-containing rocks and calcium rich sediments; low to high elevations; Greenland; Alta., B.C., N.W.T., N.S., Nunavut, Que.; Alaska, Ark., Colo., Iowa, Mass., Mich., Minn., Mont., N.H., N.Y., Tenn., Vt., Wis.; Eurasia.

The stem of Mesoptychia badensis is frequently branching proximal to the perianth. The male plants are very small, ca. 0.3--0.35 mm wide.

2. Mesoptychia bantriensis (Hooker) L. Söderström & Váňa, Phytotaxa 65: 52. 2012

Jungermannia bantriensis Hooker, Brit. Jungermann., 41. 1816; Lophozia bantriensis (Hooker) Stephani; Leiocolea bantriensis (Hooker) Jørgensen

Plants 8--18 x 1.8--2.8 mm, prostrate to loosely ascending in dense patches, brown to green-brown or yellowish brown.  Stem 245--420 \um wide, 220--390 \um high; in section outer cells strongly thick-walled, ca. 25 \um, inner cells thinner-walled, ca. 25--35 \um.  Rhizoids brownish, in tufts.  Leaves contiguous, inserted at ca. 30--40° with axis, dorsally shortly decurrent, ventrally subtransversely inserted, 845--1640 x 945--1645 \um, 0.9--1.15 times as long as wide, plane with lobes frequently incurved or flexed towards the shoot apex, trapezoidal to orbicular or broadly oval, asymmetrical, divided 1/7--1/5(--1/3) of leaf length, sinus angular or crescentic, lobes arcuate to biconcave, acute.  Cells in midleaf 37--50 x 29--45 \um, elongate, walls thin, trigones large, bulging, marginal cells 30--48 \um, thin-walled, but with thickened external wall;   cuticle coarsely papillose throughout.  Oil-bodies 2--8(--12) per cell, spherical and 4--9 \um to ellipsoidal and up 10--15 x 8 \um.  Underleaves regularly present, lanceolate, ca. 300--650 \um, frequently with acute teeth, teeth sometimes reduced to slime papillae.  Specialized asexual reproduction absent.  Sexual condition dioicous.  Androecia bracts similar to sterile leaves, base not or obscurely inflated, antheridia 2 per bract.  Perianth  tubular to tubular-subclavate, smooth or shallowly plicate, abruptly contracted to the beaked mouth,  emergent from bracts for 1/2--2/3 of the length, mouth dentate;   bracts trapezoidal, 2-lobed 1/5--1/4 of the length, sinus gibbous, lobes acute;  stem rarely branched proximal to perianth.     

Moist, sandy, calcium-rich or, at least, basic soil and rocks near streams, also wet basic tundra; 0--3200 m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., Nunavut, Que.; Alaska, Calif., Colo., Idaho, Minn., Mont., Nev., Oreg., Utah, Vt., Wash.; Eurasia.

3. Mesoptychia collaris (Nees) L. Söderström & Váňa, Phytotaxa 65: 53. 2012

Jungermannia collaris Nees, Fl. Crypt. Erlang., xv. 1817; Jungermannia muelleri Nees ex Lindenberg; Leiocolea alpestris (Schleicher) Isoviita; L. muelleri (Nees ex Lindenberg) Jørgensen; Lophozia alpestris (Schleicher) A. Evans; Leiocolea collaris (Nees) Jøgensen

Plants 5--10 x 1.6--1.9 mm, loosely ascending, brown to green brown.  Stem 250--350 \um wide, 200--250 \um high; in section outer cells ca. 17--22 \um in diameter, walls thick, median lamina often darker; inner cells gradually larger, mostly 6--7-sided, ca. 20--28 \um in diameter, walls thin to slightly thickened.  Rhizoids brownish, numerous, in indistinct fascicles.  Leaves contiguous, inserted at 15--40º with axis, 850--875 x 825--935 \um, 0.95--1.1 as long as wide, plane or slightly concave, oblique lingulate to oblique oval and oblique trapezoidal, nearly symmetrical, 2-lobed for 1/5--1/4 of the leaf length, sinus angular, U-shaped or gibbous, lobes arcuate to triangular, blunt to rarely acute.  Cells in midleaf 33--43 x 22--35 \um, walls thin, brownish, trigones distinct, concave, marginal cells 28--38 \um, walls thin, brownish, trigones triangular to slightly convex;   midleaf cuticle obscurely to evidently papillose.  Oil-bodies 2--5(--9) per cell, spherical, 4--5 \um, to ellipsoidal, to 14(--21) \um.  Underleaves regularly present to irregularly present, lanceolate, ca. 270--450  \um.  Specialized asexual reproduction absent.  Sexual condition dioicous.  Androecia intercalary, resembling sterile shoot segments, bracts similar to sterile leaves but frequently with one additional tooth near dorsal base, bracteole larger than underleaves, lanceolate,  entire; antheridia 1 per bract.  Perianth  tubular to cylindrical-clavate, smooth, abruptly contracted to the beaked mouth, emergent from bracts for 2/3--3/4 of the length, mouth crenulate to shortly ciliate;   bracts trapezoidal, 2-lobed 1/5--1/4 of the length, sinus gibbous, lobes acute.  

Wet basic tundra, over limestone on boulders along streams, also wet basic cliffs, mostly in non-forested areas; low to high elevations (0--3000 m); Greenland; Alta., B.C., N.W.T., Sask., Yukon; Alaska, Calif., Colo., Idaho, Mont., Nev., Oreg., Wash., Wyo.; Eurasia.

The male plants of Mesoptychia collaris are smaller than the female. Prior to R. Grolle (1971), the name Lophozia alpestris was misapplied to L. sudetica  (Nees ex Huebener) Grolle; virtually all references to L. alpestris in the North American literature refer to L. sudetica.

4. Mesoptychia gillmanii (Austin) L. Söderström & Váňa, Phytotaxa 65: 53. 2012

Jungermannia gillmanii Austin, Bull. Torrey Bot. Club 2(3): 12. 1872; Leiocolea kaurinii (Limpricht) Jørgensen; Lophozia gillmanii (Austin) R. M. Schuster; L. gillmanii var. ciliolata R. M. Schuster; Leiocolea gillmanii (Austin) A. Evans

Plants 7--20 x 1.75--2.45 mm, prostrate, brownish green to green-brown.  Stem 175--280 \um wide, 160--270 \um high; in section cells ca. 25--38 \um, surface crenulate, outer cell walls thickened (particularly the external wall),  inner cell walls thinner.  Rhizoids brownish, in dense  tufts.  Leaves distant to contiguous, inserted at ca. 15--20° with axis, dorsally decurrent in the biggest shoots, ventrally subtransversely inserted, 950--1000 x 1000--1050 \um, 1--1.1 as long as wide, plane to slightly convex, obliquely trapezoidal, symmetrical to asymmetrical, 2-lobed for 1/9--1/5 of leaf length, sinus angular or crescentic, lobes triangular to arcuate, usually narrowed to an acute tip.  Cells in midleaf 35--50 x 25--33 \um, elongate, walls thin, colorless to grayish, trigones concave, marginal cells 20--28 \um, trigones moderate in size, concave to loosely convex; cuticle papillose throughout.  Oil-bodies 2--8(--9) per cell, spherical, (3.5--)5--8(--10) \um in diameter to ellipsoidal, to 12--13 \um.  Underleaves irregularly present, lanceolate, ca. 450--700  \um, commonly with a few teeth on each side.  Specialized asexual reproduction absent.  Sexual condition paroicous.  Androecia in 2--3(--4) pairs of bracts, separated from perichaetium by 1 pair of sterile leaves, bracts obliquely trapezoidal, frequently with obtuse teeth on dorsal side.  Perianth  tubular to ellipsoidal, smooth, suddenly tightened to the beaked mouth,  emergent from bracts for 2/3 of the length, mouth dentate;   bracts ovate to obliquely ovate,  sometimes with teeth in middle of dorsal side.   

Wet moist calcium rich rocks, cliffs near waterfalls, calcareous solifluction spots near snow-beds, basic swamps; low to high elevations (0--3000m); Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., Nunavut, Ont., Que., Yukon; Alaska, Ariz., Calif., Idaho, Maine, Mass., Minn., Mont., N.H., N.Y., Ohio, Oreg., Vt., Wash., Wis., Wyo.; Eurasia.

The bracteoles of Mesoptychia gillmanii are small, 2-fid, with lobes of different sizes. The spores are ca. 15 \um, finely papillose, and the elaters are ca. 8 \um in width.

5. Mesoptychia heterocolpos (Thedenius) L. Söderström & Váňa Phytotaxa 65: 53. 2012

Jungermannia heterocolpos Thedenius, Kongl. Vetensk. Acad. Handl. 1837: 52. 1838; Leiocolea holmeniana (Inoue & Steere) Konstantinova; Lophozia heterocolpos (Thedenius) M. Howe; Leiocolea heterocolpos (Thedenius) H. Buch

Plants 5--20 x 0.9--1.3 mm, prostrate to ascending, with erect shoot apices bearing gemmae, green, apices deep green to brownish deep green, older shoots brown.  Stem ca. 250 \um wide, 190--240 \um high; in section outer cells with thick walls, ca. 12--17 \um in diameter, inner cell walls thin to slightly thickened, ca. 22--28 \um in diameter.  Rhizoids brownish to almost colorless, sparse, in tufts.  Leaves distant to contiguous, inserted at ca. 35--45° with axis, 475--650 x 525--625 \um, 0.1--1.1 as long as wide, plane to slightly concave, ovate to trapezoidal, nearly symmetrical, 2-lobed for 1/5-1/4 of the length, sinus angular or U-shaped, lobes arcuate.  Cells in midleaf 25--33 x 19--22(--30) \um, thin-walled, trigones large and bulging, marginal cells 14--22 /mu;   cuticle mostly virtually smooth, rarely papillose.  Oil-bodies 2--8(--12) per cell, spherical, 5--8(--11) \um, to ellipsoidal or tubular, 6--14(--16) x 4--7(--10) \um.  Underleaves regularly present, 2-fid, divided almost to the base (lobes connected by only 1--2 cells at base), ca. 200--350 \um.  Specialized asexual reproduction by gemmae present on leaf apices of modified erect isophyllous shoots or unmodified branch apices, leaves bearing gemmae of elongate cells with small trigones; gemmae brown, 25--30 x 15--20 \um, 2-celled, ellipsoidal.  Sexual condition dioicous.  Androecia intercalary, semi-spicate, bracts in 3--4 pairs, concave, inflated at base, sometimes with deflexed lobe apices, trapezoidal to ovate, with additional teeth on dorsal side, antheridia 2 per bract.  Perianth   clavate to oblong ovate, suddenly or gradually narowed to the beacked mouth, emergent from bracts for 2/3--3/4 of the length, mouth dentate;   bracts trapezoidal, 2-lobed 1/5--1/3 of the length, sinus gibbous, lobes acute. 

Varieties 3 (2 in the flora): North America, Eurasia.

The elaters of Mesoptychia heterocolpos are 7--8 \um in width, and the spores 11--13 \um, finely papillose.

1. Leaves plane to weakly concave, inserted mostly at 35--40º with axis, gemmiparous shoots common …5a. Mesoptychia heterocolpos var. heterocolpos

1. Leaves distinctly concave, inserted mostly at 40--50º with axis, gemmiparous shoots rare … 5b. Mesoptychia heterocolpos var. harpanthoides

5a. Mesoptychia heterocolpos var. heterocolpos

Plants ascending to semi-erect in dense moss patches. Leaves plane to weakly concave, inserted mostly at 35--40º with axis. Cells in midleaf 18--28 \um wide. Gemmiparous shoots common.

Wet or moderately dry basic or neutral cliffs covered with humus, patches in oligotrophic bogs, rarely on decaying wood; 0--3350m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., Nunavut, Ont., Sask., Yukon; Alaska, Calif., Colo., Idaho, Maine, Mich., Minn., Mont., Nev., N.H., N.C., Oreg., Utah, Vt., Wash., Wis., Wyo.; Eurasia.

A specimen from Ohio, as noted by Schuster (1969), requires verification.

5b. Mesoptychia heterocolpos var. harpanthoides (Bryhn & Kaalaas) L. Söderström & Váňa, Phytotaxa 65: 53. 2012

Lophozia harpanthoides Bryhn & Kaalaas, Rep. Second Norweg. Arctic Exped. Fram 1898---1902, 11: 31 1906; Lophozia heterocolpos var. harpanthoides (Bryhn & Kaalaas) R. M. Schuster; Leiocolea harpanthoides (Bryhn & Kaalaas) A. Evans; L. heterocolpos var. harpanthoides (Bryhn & Kaalaas) S. W. Arnell

Plants commonly erect, growing in dense moss patches. Leaves clearly concave, inserted mostly at 40--50º to stem. Cells in midleaf 18--28 \um wide. Gemmiparous shoots rare.

Calcareous to neutral mossy tundras, as a rule among mosses in dense patches, and near snow-beds; low to moderate elevations; Greenland; Nunavut, Que.; n Eurasia.

6. Mesoptychia polymorpha Stotler, Crandall-Stotler & Bakalin, Polish Botanical Journal 58: 82. 2013

Plants 3--6 x 0.6--1.5 mm, prostrate to ascending, pale yellow green without secondary pigments.  Stem 140–180 \um wide and 100–125 \um high, in section cells more or less uniform, with thin walls and lacking trigones, 25--35 \um in diameter, external wall of stem slightly thicker.  Rhizoids sparse to abundant, grayish brown.  Leaves contiguous to distant, highly variable in shape, from broadly obovate to obtrapezoidal, rectangular or lingulate, with the apices varying from obtuse to truncate, or retuse, emarginate, with the lobes mostly obtuse and unequal in size, inserted at ca. 15--40° with axis, dorsally not decurrent, ventrally subtransversely inserted, 250--700 x 250--600 \um, 0.9--1.1 times as long as wide, plane.  Cells in midleaf 30--65 x 27--46 \um, walls thin, trigones absent to vestigial, marginal cells 27--45 \um, thin-walled;   cuticle finely striolate.  Oil-bodies (2--)3--5(--8) per cell, elliptical to spheroidal, gray, coarsely granulate, 3.8–5.7 µm in diameter and up to 9.5 µm long.  Underleaves irregularly present and easily deciduous, mostly 2--6(--8) cells long, ca. 40--280 \um.  Specialized asexual reproduction absent.  Sexual condition paroicous.  Androecia intercalary, bracts in 2--4 pairs,  concave to cupped and channeled, commonly with one tooth near dorsal leaf base, antheridia 1(--2) per bract, bracteole absent or highly reduced.  Perianth clavate to obpyriform, smooth, suddenly tightened to the beaked mouth,  emergent from bracts for 1/2--2/3 of its length, mouth crenulate to dentate; bracts  trapezoidal,  2-lobed for 1/6--1/5 of the length, sinus gibbous, bracteole absent or up to 120 \um long. 

Continuously moist, deeply shaded calcareous substrates; 45--210 m; Calif.

The stem of Mesoptychia polymorpha is rarely branched, with branches only of the lateral, endogenous Plagiochila-type.

7. Mesoptychia rutheana (Limpricht) L. Söderström & Váňa Phytotaxa 65: 54. 2012

Jungermannia rutheana Limpricht, Jahresber. Schles. Ges. Vaterl. Cult. 61: 207 1884: Lophozia rutheana (Limpricht) M. Howe; Leiocolea rutheana (Limpricht) Müller

Plants 30--150 x 4.2--4.9 mm, prostrate, brown to greenish brown.  Stem ca. 400 \um wide, 290 \um high, in section outer cells 15-30 \um in diameter with thick walls, inner cell walls thin to slightly thickened.  Rhizoids dense to sparse or virtually absent, short, brownish to almost colorless, isolated or in tufts.  Leaves contiguous, inserted at ca. 10--15° with stem, dorsally barely decurrent, 1625--2520 x 1925--3100 \um, 0.7--0.85 as long as wide, slightly convex to almost plane, obliquely trapezoidal, asymmetrical, widest at middle or just proximal, 2-lobed for ca. 1/5 of length, sinus gibbous to crescentic, frequently valvate in margin near sinus base, lobes angular, arcuate to biconcave, with blunt to acute apices.  Cells in midleaf 43--60 x 30--38 \um, thin-walled, walls brown, trigones large and bulging, marginal cells 22-30 \um in diameter, with thin to slightly thickened walls and convex trigones;   cuticle coarsely papillose.  Oil-bodies 2--6 per cell, mostly ellipsoidal, 7--20 x 5--9 \um.  Underleaves regularly present, 1--3--lobed, middle lobe commonly larger, ca. 1400--1750 \um (lateral laminae ca. 1000 \um), commonly with 2--10 laciniae and cilia spreading at narrow angles.  Specialized asexual reproduction absent.  Sexual condition paroicous.  Androecia proximal to perichaetium, in 1--2 pairs of bracts, similar to sterile leaves but loosely inflated in basal part, antheridial stalk 2--seriate.  Perianth  fusiform, loosely folded, gradually narrowed to the mouth, emergent from bracts for 3/4--4/5 of length, mouth not beaked;   bracts just proximal to perianth, in 1 pair, smaller than sterile leaves, 2-lobed for 1/5 of length, sinus narrowly gibbous or angular, acute lobes.   

Rich fens, springy swamps, moist mossy tundras, exclusively on basic substrates; 0--1300 m; Greenland; Alta., B.C., Man., Nfld. and Labr. (Nfld.), N.W.T., Que., Yukon; Alaska, Minn., Mont., N.Y.; Eurasia.

The elaters of Mesoptychia rutheana are 8--10 \um wide and the spores are (12--)16--20 \um, pale brown.

8. Mesoptychia sahlbergii (Lindberg & Arnell) A. Evans, Ottawa Naturalist 17: 15. 1903

Jungermannia sahlbergii Linsberg & Arnell, Kongl. Svenska Vetensk. Acad. Handl., n.s. 23(5): 40. 1889; Lophozia sahlbergii (Lindberg & Arnell) Stephani

Plants 30--60(--90) x 3--5 mm, prostrate, brownish red to red-purple, rarely brown.  Stem 300--500 \um wide, 250-400 \um high, in section outer cells 15-30 \um with thick walls, inner cell walls thin to slightly thickened.  Rhizoids dense, short to long, brownish to almost colorless, forming mat under stem.  Leaves contiguous to subimbricate, inserted at ca. 10--15° with stem, dorsally barely decurrent, 1400--3000 x 1500--3500 \um, 0.7--0.9 as long as wide, slightly convex to almost plane, loosely plicate along midline, obliquely trapezoidal, asymmetrical, widest near base, 2-lobed for 1/10--1/5 of length, sinus gibbous to crescentic, frequently valvate in margin near sinus base, lobes angular, mostly arcuate, with blunt to acute apices.  Cells in midleaf 30--50 x 32--40 \um, thin-walled, walls brown, trigones large and bulging, marginal cells 22--30 \um in diameter, with thin to slightly thickened walls and convex trigones; cuticle coarsely papillose.  Oil-bodies 3--8 per cell, mostly ellipsoidal, rarer nearly spheric, 6--12 x 4--6 \um.  Underleaves regularly present, 1--2--lobed, 1000--1800 \um, with numerous cilia spreading at right angles.  Specialized asexual reproduction absent.  Sexual condition dioicous.  Androecia, in 2--4 pairs of bracts, similar to sterile leaves but loosely inflated in basal part and with additional small lobe at dorsal side, antheridial stalk 2-seriate.  Perianth  fusiform, loosely folded, gradually narrowed to the mouth, emergent from bracts for 3/4--4/5 of length, mouth not beaked; perigynium well developed, ca. the same length as perianth, at right angle with stem, strongly rhizogenous, bracts just proximal to perianth, in 1 pair, larger than sterile leaves, 2-lobed for 1/5 of length, sinus narrowly gibbous or angular, lobes acute.   

Rich fens, springy swamps, moist mossy tundras, exclusively on basic substrates; 0--500 m; Greenland; Nunavut; Yukon; Alaska; n Eurasia.

The elaters of Mesoptychia sahlbergii are 7--9 \um wide and the spores are 12--15 \um, brown.

5. RIVULARIELLA D. H. Wagner, Phytoneuron 2013-10: 2. [Lat. rivularis, of brooklets, and ella, diminutive, alluding to habitat]

David H. Wagner

Plants medium-sized, dark. Stems without internal differentiation, little branched, rhizoids scattered on ventral surface. Leaves succubous, emarginate to shallowly lobed, insertion from nearly horizontal to transverse, oil bodies present in all leaf cells. Underleaves small to large, most prominent on erect shoots. Asexual reproduction by fasciculate gemmae produced on leaf margins. Sexual condition monoicous, paroicous or heteroicous. Androecia with 2--3 antheridia per bract, stalks 1-seriate.  Gynoecia terminal on main shoot, bracts and bracteoles larger than leaves, perianth oblong to fusiform, plicate. Sporophyte seta massive; capsule wall several layers thick, outer layer with nodular thickenings, inner layers with semiannular thickenings. Spores greenish when fresh. Elaters mostly 2(--3)-spiral.

Species 1 (1 in the flora): w,nw North America.

Rivulariella is a monotypic genus of the Northwest. It is found attached to pebbles in the bottom of small streams.

SELECTED REFERENCE  Evans, A.  1938.  A new species of Chiloscyphus from Utah.  Bryologist 41: 50--57.

1. Rivulariella gemmipara (A. Evans) D. H. Wagner, Phytoneuron 2013-10: 2. 2013  E F

Chiloscyphus gemmiparus A. Evans. Bryologist 41: 50. 1938

Plants medium sized, shoots to 30 mm, to 4 mm wide, prostrate to ascending, dark green, blackish at base.  Stems slightly to strongly dorsiventrally compressed; prostrate stems flattened, 300--500 \um wide, 150--200 \um high tightly adherent to substrate; stems of ascending shoots more nearly cylindrical, to 400 \um in diameter; branches mainly lateral-intercalary on prostrate shoots, often abundant; branches sparse on ascending shoots, mainly terminal; cortical cell walls thin, 15 x 29 \um, medullary cells 30 x 40 \um with slightly thicker cell walls; rhizoids scattered, abundant on prostrate stems, absent on ascending stems.  Lateral leaves variable 120--150 x 150--200 mm, inserted obliquely and spreading on prostrate shoots, nearly transverse, imbricate and concave on ascending shoots, obovate, rounded or truncate to emarginate or shallowly bilobed.  Median cells subquadrate to elongate-rectangular, rarely hexagonal, 16--18 x 17--25 \um, cell walls thin, trigones absent or poorly developed, marginal cells slightly smaller, cuticle smooth.  Oil-bodies (5--)7--12(--20) per median leaf cell, ovoid, 4 x 6 \um, finely granular, colorless.  Underleaves absent or fugacious on prostrate stems, usually small, few celled and multilobed or subulate to lanceolate on ascending stems, with larger underleaves occurring sporadically and randomly, occasionally nearly the same size and shape as the lateral leaves on erect shoots, these shoots virtually isophyllous.  Asexual reproduction by fasciculate gemmae produced along leaf margins at shoot tips, ovoid at maturity, 120--180 x 160--300 \um, composed of 2--4(--6) cells each.  Sexual condition paroicous or heteroicous.  Androecia either below gynoecia or on separate shoots, antheridia in bracteoles of isophyllous shoots on paroicous shoots or in bracts little different from vegetative leaves on anisophyllous shoots when strictly male, spicate or not.  Gynoecia on erect, isophyllous shoots, archegonia developing before perianth formation, perianth exerted 1/2 or more above bracts, inflated, oblong to fusiform, smooth and cylindrical proximally, 4--5 plicate distally, narrowed to an entire or weakly crenulate mouth; perigynium absent. 

Alpine or subalpine zones, strictly confined to small streams fed by cold water springs, attached small stones or pebbles in the stream bed, best developed in full sun; Alaska (Unalaska Island), California, Oregon, Utah, Washington.

The gemmae of Rivulariella gemmipara are abundant in some populations but nearly absent in others. The spores are noticeably green when fresh.  The capsule wall is several cell layers thick, only the outer layer having nodular thickening and the inner cells with semiannular thickenings. Other liverworts commonly found growing with Rivulariella are Jungermannia eucordifolia, Chiloscyphus polyanthos, and Scapania undulata.  Two lichens are also found in this association: Peltigera hydrothyria Miadl. & Lutzoni and Leptogium rivale Tuck.