BFNA Title: Jungermannia
Author: V. Bakalin
Date: September 2, 2011
Edit Level: R
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
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XX. 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; 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) 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


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.