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BFNA Title: Jungermannia |
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XX.
JUNGERMANNIA Linnaeus, 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, 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 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., 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; 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.), 6. Jungermannia
pumila Withering,
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; 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. |
