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Scapaniaceae – Diplophyllum
XXX.
DIPLOPHYLLUM Dumortier, Syll. Jungerm. Eur. P. 44,
1871. Recueil
d'Observations sur les Jungermanniacées 15. 1835
* [Latin diplo double, phyllum leaf, alluding to the folded leaf]
Jungermannia sect. Diplophyllum Dumortier, Syll. Jungerm. Eur. 44. 1831
Paul L. Redfearn, Jr.
Plants usually in thin flat patches
or mats. Stems with few intercalary and axillary branches, usually
with subfloral innovations; cortex in 3--5 layers, cells somewhat to strongly
flattened, smaller, more strongly thick-walled than medullary cells; branches
few; rhizoids scattered, colorless. Leaves complicate-2-lobed with
smaller dorsal and larger ventral lobes, not decurrent, the most
proximal leaves usually smaller, rounded to apiculate, fold 1/4--1/2 the
length of leaf, sheathing, nearly or almost parallel to the stem, distal half
gradually arching outward 45--100º and becoming distinctly folded but not winged; ventral half of leaf
narrowly lingulate, ± falcate, 2--3.5:1; dorsal half similar but smaller and
shorter; lobes strongly divergent, the dorsal 0--45º angle, narrowly obovate
to lingulate; ventral lobes usually divergent at a 60--110º ; leaf margins
entire to more commonly finely crenulate to denticulate with small, sharp
1-celled teeth;
cell
walls little thickened at corners, marginal cells in several rows of
strongly, equally thick-walled; medial cells with small or inconspicuous
trigones with slightly thickened or non-thickened walls, usually with
conspicuous papillae, appearing nearly opaque compared to marginal cells;
cells of the sheathing base elongate, 3--5:1; oil bodies distinctly
segmented, appearing papillose, often absent in marginal cells, 2--5 per intramarginal cell, 10--16 or more per elongated cell. Underleaves
absent. Specialized asexual reproduction by 1--2-celled, stellate
gemmae with strongly protuberant angles.
Sexual condition dioicous, rarely autoicous or paroicous. Androecia
with bracts imbricate in 4--8 pairs, similar to leaves, antheridia 1--3
per bract. Gynoecia terminal,
bracts imbricate, similar to leaves; perianth ovate
to cylindric, ± plicate, terete to slightly flattened near apex, apex
denticulate, sometimes lobed or laciniate. Sporophyte capsule exserted on a
short seta, ovoid, 3--5-stratose. Spores
11--15 \um, surface with irregular network of fine vermiculate, partly
anastomosing markings, elevated at intersections as ±
weak tubercles; elaters 2-spiraled.
Species
24 (6 in
the flora); North America, South America, Europe, Asia, Africa,
Australia (Tasmania).
Diplophyllum is recognized by conduplicate, non-decurrent leaves with a defined
marginal border, ventral lobes rounded-ovate, larger than dorsal lobes,
underleaves absent; perianth cylindric and distally
pluriplicate (furrowed in subgenus Macrodiplophyllum, which has been transferred to Scapania). Diplophyllum plicatum is
placed in the genus Scapania by A. D. Potemkin (1999).
SELECTED
REFERENCES: Potemkin, A. D. 1999. Circumscription of the family Scapaniaceae, with segregation of the new family Diplophyllaceae (Hepaticae).
Annales Botanici Fennici
36: 271--283. Schuster, R. M. 1975. The Hepaticae
and Anthocerotae of North America east of the Hundredth Meridian 3: 177--232.
1.
Leaves with margins entire; shoots 3--5(--8) mm, dry leaves slightly to
strongly dorsally secund and connivent; perianth barely dorsiventrally
flattened, gradually narrowed to a pointed apex . . . ..6. Diplophyllum andrewsii
1.
Leaves with margins distinctly crenulate or denticulate with projecting
1-celled teeth; shoots 5--25 mm, dry leaves slightly dorsally secund; perianth, ±
strongly dorsiventrally compressed, little to
strongly contracted to mouth.
2. Leaf lobes with a sharply
defined vitta of extremely elongate (60--70
\um) linear cells; cuticle often smooth
or virtually so; cortex 3--5-stratose . . . . .1. Diplophyllum albicans
2. Leaf lobes without a
distinctly defined vitta of strongly elongated
cells; cuticle distinctly papillose; cortex in 1--2 layers.
3. Ventral leaf lobes
distinctly apiculate . . . 3. Diplophyllum apiculatum
3. Ventral leaf lobes rounded
at apex, usually broadly so.
4. Dioicous, often sterile and gemmiparous; medial and submedial
cells (distal to keel) of lobes short-rectangular, ca. 1.5--2:1; cells of
apex of ventral lobes 7--10.5 \um; gemmae at maturity greenish,
1--2-celled;. . . .2. Diplophyllum taxifolium
4. Autocious
or paroicous, perianths nearly always present; medial and submedial
cells of ventral (and usually dorsal lobes) lobes sublinear,
2--4:1; cells of apex of ventral lobes 8--12 \um; gemmae brownish at maturity
(or absent).
5. Ventral lobes never
truncate, not parallel-sided, distally broadly rounded; dorsal lobes often
obtuse or subacute; medial leaf cells coarsely and closely papillose; perianth mouth with terminal cells not or barely
elongate.. 4. Diplophyllum obtusatum
5. Ventral leaf lobes often
truncate at apex, nearly or quite parallel-sided; medial leaf cells usually weakly papillose; perianth mouth with some terminal cells strongly elongate
. . .. . . . 5. Diplophyllum obtusifolium
1. Diplophyllum albicans (Linnaeus) Dumortier, Recueil
d'Observations sur
les Jungermanniacées,
16. 1835
Jungermannia albicans Linnaeus, Sp. Pl., 1133. 1753
Plants in compact to loose mats with
crowded erect branches, 2--5 cm, green, golden yellow or deep brown, Arctic
plants sometimes scorched, older portions frequently green, distal portions
often reddish. Stems brownish,
rigid, 1--5 cm, 0.2--0.3 mm in
diameter, branches sparse, usually erect and simple; cortex 3--5 layers
thick; leafy shoots 1.5--3.5
mm wide; rhizoids few.
Leaves distinctly close together to imbricate, strongly
complicate-2-lobed, strongly dorsally secund and connivent
when dry, laterally spreading when moist, lobes with distinctly defined vitta of 4--6 rows of glistening, extremely elongated
cells, 60--70 x 12--16 \um, leaf margins sparingly to strongly denticulate to
serrate; marginal cells equally thick-walled, 13--22 x 12--14 \um , medial
cells less thick-walled, 13--22 x 12--16
\um; cuticle smooth to almost imperceptibly verruculose; dorsal lobes
oblong-ovate to lingulate, apex obtuse to subacute, 0.5--1.5 x 0.25--0.45 mm,
ca. 0.5 size of ventral lobe, erect; ventral lobes similar to dorsal lobes,
very narrowly lingulate to oblong-ovate, rounded, obtuse to subacute at apex,
1--2 x 0.35--0.45 mm,
widely spreading, frequently falcate, free lobe at 75--100º angle with
stem. Specialized asexual
reproduction by abundant gemmae, usually restricted to excurrent apices
of vittae, 1-celled, stellate, usually yellowish
green, becoming fuscous in scorched Arctic plants. Sexual condition dioicous. Androecia short, bracts 4--8
pairs, often reddish, antheridia 1--2 per bract. Gynoecia bracts similar to leaves
but somewhat larger; perianth half exserted at
maturity, distally narrowed and plicate, mouth lobes lobulate, ending in
cilia bearing 1 to several 1-celled teeth at base. Sporophyte capsule ovoid. Spores 11--15 \um, papillose,
yellowish brown.
Shaded,
humid logs, tree trunks, noncalcareous rock walls;
0--500 m.
Greenland; B.C., N.B., Nfld. and Labr.
(Nfld.), N.S., Nunavut, Yukon;
Alaska, Oreg.,
Maine, Wash.;
Europe; Asia; Atlantic
Islands.
Diplophyllum
albicans has
an oceanic distribution in the coniferous biome north to the Arctic tundra.
2. Diplophllum taxifolium
(Wahlenberg) Dumortier, Recueil
d'Observations sur les
Jungermanniacées, 16. 1835
Jungermannia taxifolia Wahlenberg, Flora Lapponica,
389. 1812
Plants
in thin,
extensive patches, green to yellowish brown.
Stems 1--3.5 cm, 0.16--0.3 mm in diameter, prostrate at base,
ascending distally; slightly branched
below perianth; cortex 1--2 layers thick;
leafy shoots 1.3--2.4 mm
wide; rhizoids few. Leaves equal in size, close together to subimbricate, lobes 0.33--0.5:1, dry leaves ± dorsally
secund; dorsal lobe appressed to stem, spreading laterally at an angle of
10--40 º, elliptic, 0.43--0.65 x 0.18--0.35 mm, male shoots
0.6--0.8 x 0.26--0.36 mm on female shoots, tapering distally to a bluntly
obtuse or somewhat rounded apex, rarely acute, varying from nearly entire to
denticulate, dorsal base closely and sharply crenulate-denticulate in distal
fifth by projecting cells, ca. 1/2 as long as ventral lobe; ventral lobe
horizontally spreading, nearly flat, somewhat falcate, distal half spreading
at an angle of 65--110º on mature leaves, lingulate-elliptic, parallel-sided
for about the medial third, somewhat tapering in the apical fifth to a
broadly rounded, bluntly obtuse apex, often more pointed on male and gemmiparous plants, 0.75--0.9 x 0.28--0.36 mm on males
shoots, female shoots 1--1.2
cm, 0.35--0.45 mm wide, margins with scattered
denticulations on apical portion, sometimes nearly entire, basal half
distinctly sharply crenulate by projecting cells; keel suberect, gradually
spreading outward; marginal cells of ventral lobe 7--10 \um near apex, mostly
isodiametric, equally thick-walled, forming a slightly to obscure pachydermous border; cells on middle ventral half ca. 1.5--2:1, 18--25 x 10--15 \um,
occasionally longer, trigones none or minute; cuticle moderately papillose,
papillae often extending onto marginal cells, 6--12 per cell in leaf middle;
oil bodies absent in 1--2 marginal cell rows, 2--3 in the next rows to 3--8 in cells of leaf middle to
10--15 in
elongated basal cells. Specialized asexual reproduction common, as masses of 2-celled gemmae at apex
of shoots, on lobe margins of distalmost leaves,
yellowish green, polygonal and stellate because of blunt projecting angles, gemmiparous leaves often twice the normal size. Sexual condition dioicous, largely
sterile, male and female plants usually in separate patches, distinctly
heterothallic, male plants smaller. Androecia
with bracts in 5--6 or
more pairs, similar to normal leaves, but ventral with lobe often more
tapering and pointed, dorsal lobe covered with wart-like papillae, antheridia
usually 2 per bract, 1 often aborted. Gynoecia
terminal, bracts slightly larger than stem leaves; perianth
0.5--0.75 exserted at maturity, ovate, dorsiventrally
compressed, plicate in distal half, a dorsal sulcus extending to near base,
strongly contracted to mouth, lobed, dentate, hyaline, occasionally
entire. Sporophyte capsule ovoid, brown. Spores 13--15 \um, closely coarsely
verruculose, pale brown.
Varieties 3
(3 in the flora): widely distributed in the northern hemisphere.
1.
Ventral lobes abruptly mucronate with a 1-celled mucro, or triangularly
narrowed and then mucronate. . . . . .2c. Diplophyllum taxifolium var.
mucronatum
1. Ventral lobes broadly
rounded.
2. Cuticle covered with many
small cuticular papillae, leaves not very opaque. . . . . . . . . . . . .2a. Diplophyllum taxifolium var. taxifolium
2.
Cuticle medially on the leaf with coarse, dense papillae, leaves very opaque
except on margins. . . . . . .2b. Diplophyllum taxifolium var. macrosticta
2a. Diplophyllum taxifolium var. taxifolium
Ventral
lobes rounded; cuticle with small, numerous papillae; leaves not very opaque.
Shaded
rocks, cliffs, soil banks, humus, often along streams; 0--1950 m; Greenland; B.C.,
N.B., Nfld. and Labr., N.S., Ont.; Alaska, Conn., Maine, Mass., Minn., N.H.,
N.Y., N.C., Tenn., Vt., Wash., W.Va.; Europe; Asia.
2b. Diplophyllum taxifolium var. macrosticta H. Buch, Commentat.Observ. Jungerm. 3(1): 23. 1928
Ventral
lobes rounded, papillae dense, cells very opaque except marginally.
Moist,
shaded rock walls and ledges, soil; 1860--2010 m; N.S.; Alaska, Maine, Minn., Tenn., Vt.; Europe; Asia.
2c. Diplophyllum taxifolium var. mucronatum R. M.
Schuster, Hepaticae
and Anthocerotae of North America, East of the Hundredth Meridian 3: 203,
plate 334. 1974 E
Leaves
abruptly mucronate with a 1-celled mucro, or triangularly narrowed and then
mucronate, cells rather opaque.
Shaded
rocks; 1645 m;
N.C.
3. Diplophyllum apiculatum
(A. Evans) Stephani, Spec. Hep. 4:110, 1910 E
Diplophylleia
apiculatum A. Evans, Bot. Gaz. 34: 372, plate 12. 1902
Plants
in patches,
green in shade to brownish or reddish in sun.
Stems 0.3--1 cm,
0.15--0.2 mm in diameter, prostrate, apices weakly ascending, sparing
branched; cortex poorly defined,1--2 layers thick; leafy shoots 1.5--2.4 mm wide; rhizoids
numerous below. Leaves nearly
equal in size along the entire stem, imbricate to subimbricate;
dorsal lobe diverging at an angle of 10--40º, appressed, elliptic, 0.23--0.48
x 0.13--0.23 mm, ca. 1/2 size of ventral lobe, apex triangularly narrowed and
usually strongly apiculate, margins irregularly denticulate, rarely entire;
ventral lobe horizontal, sometimes weakly dorsally secund when dry, ± falcate
and gradually spreading, apical half spreading 65--105º, narrow and
lingulate-pointed, with nearly parallel sides, 0.5--0.75 x 0.2--0.3 mm, averaging 2.5:1,
apex triangularly narrowed, usually strongly apiculate, terminated by a tooth
often 2 cells long, margins ± sparingly denticulate toward apex, distinctly
crenulate along basal half of ventral margin; marginal cells of apex nearly
isodiametric, 12--16 x 9--14 \um, strongly thick-walled in 3--5
rows, inner cells gradually thin-walled, not collenchymatous, 16--23 x
12--14 \um; cuticle smooth on margins,
but slightly to distinctly minutely papillose in medial parts
of lobes, papillae oval, mostly 6--12 per cell in central part of lobe; oil
bodies of marginal and submarginal cells small,
3--6 per cell, 10--20 in
medial, on submedial and ± elongate cells, almost
filling the lumen making the cells very opaque. Specialized asexual reproduction common,
gemmae in dense clusters, on apical parts of lobes of distalmost
leaves, 1--2-celled, polygonal with rounded protuberances, ± stellate in
outline, always green. Sexual
condition autoicous. Androecia usually
on main shoots, bracts 2--4 pairs, similar to leaves, strongly
inflated proximally, antheridia 1 per bract.
Gynoecia usually on short lateral intercalary branches,
originating below male shoots, bracts similar but larger than normal leaves,
5 pairs, distal 2--3 pairs suddenly larger than lower pairs; perianth 1/2 or more exserted, obovoid, somewhat dorsally
flattened, apical half 4--6-plicate, gradually narrowed toward an irregularly
lobed mouth, lobes dentate-laciniate with teeth
1--3 cells long. Sporophyte
capsule ovoid, red--brown. Spores
12--14 \um, minutely verruculose, yellow-brown.
Two
varieties are recognized by R. M. Schuster (1974).
Varieties
3 (2 in the flora): temperate North America and eastern Asia.
1.
Ventral leaf lobes ± tapering, mostly acute to apiculate, usually gradually
so, medial leaf cells with small to moderate-sized papillae, cell outlines
clearly distinguished; marginal cells prominently thick-walled, forming a
vague border; lowland form . . . ... . . 3a. Diplophyllum apiculatum var.
apiculatum
1.
Ventral lobes rounded, ± suddenly (rarely gradually) apiculate to mucronate
medial leaf cells with 2--6 coarse, juxtaposed papillae, virtually hiding the
cell outlines; marginal cells scarcely thick-walled, not forming a border;
upland form . .. . . . 3b. . Diplophyllum apiculatum var. taxifolioides
3a. Diplophyllum apiculatum var. apiculatum E
Ventral
lobes ± tapering, mostly acute to
apiculate, papillae of leaf cells delicate, medial leaf cells prominently
clearly distinguished, marginal leaf cells prominently thick-walled.
Moist
shaded soil banks, acidic rocks, beneath ledges, shaded vertical exposures,
rarely on rotting logs; 120--1400
m; Ont.; Ala., Ark., Conn., D.C., Ga., Ind., Kans.,
Ky., Maine, Md., Mass., Mich., Minn., Miss., Mo., N.J., N.Y., N.C., Ohio,
Pa., R.I., S.C., Tenn., Vt., Va., W.Va., Wis.; Asia (Japan).
Diplophyllum
apiculatum var. apiculatum is
endemic to the deciduous forests of eastern North
America.
3b. Diplophyllum apiculatum var. taxifolioides
R. M. Schuster, Hepaticae
and Anthocerotae of North America, East of the Hundredth Meridian 3: 214,
plate 338, figs. 1, 3, 8--10, 12, plate 340, figs. 6--15. 1974 E
Ventral
lobes rounded, usually suddenly apiculate, leaf cells with 2--5 coarse,
juxtaposed papillae virtually hiding the cell outlines.
Shaded
humus of damp ledges; 1860--1920
m; N. C., Tenn.
R.
M. Schuster (1974) suggested the possibility that this variety is a hybrid
between D. apiculatum and D. taxifolium.
4. Diplophyllum
obtusatum
(R. M. Schuster) R. M. Schuster, Hepaticae and
Anthocerotae of North
America, East of the Hundredth 3: 215. 1974
Diplophyllum
apiculatum var. obtusatum R. M.
Schuster, Amer. Midl. Nat. 49: 432. 1953
Plants in small, depressed mats or
patches or creeping among other bryophytes, green in shade to purplish or
reddish brown in sun, proximal leaf bases tending to become vinaceous even when plants otherwise green. Stems 0.5--1.2 cm, 0.16--0.26 mm in diameter; prostrate with ascending tips, branching on
one or both sides, or from below or axils female bracts or from older
portions of stems; cortex 2-stratose;
leafy shoots 0.8--2 mm
wide; rhizoids scattered, frequent, long. Leaves subimbricate,
often closely so in androecial region, weakly to moderately secund when dry;
ventral lobes lingulate, free distal half diverging 55--95º with stem, not
parallel-sided and widest before apex, distally broadly rounded, often obtuse
or subacute, 0.69--1.1 x 0.36--0.45 mm, 3--4 /x as long as fold; medial and submedial cells sublinear,
2--4:1, cells of apex 8--12 \um, margins distinctly denticulate except above
junction with keel, rarely subentire, proximal margins
strongly denticulate with sharp, jagged, often curved 1-celled teeth; dorsal
lobe obliquely or erect spreading, 15--55º, appressed, elliptic, obovate to
short-lingulate, usually rounded at apex, occasionally apiculate to subacute,
0.42--0.46 x 0.22--0.26 mm,
margins ± finely denticulate, keel weakly arched; cells at apex and margins
10--12 \um, medial and submedial cells sublinear, 2--4:1, 11--14 x15--18 \um, in proximal part
of leaf forming a weakly defined “vein,” opaque because of numerous large oil-bodies
and chloroplasts; marginal and submarginal cells
strongly and evenly thick-walled, inner cells often thinner walled with
minute trigones; cuticle of medial coarsely and closely papillose; oil bodies
absent in most marginal cells, 3--6 in medial cells. Specialized asexual reproduction by
numerous, mostly 1-celled gemmae, spheric to ovoid,
thick-walled, green becoming brownish at maturity, approximately stellate in
outline. Sexual condition monoicous,
usually autoicous, very rarely paroicous. Androecia bracts in 2--5 pairs,
similar but smaller than stem leaves, more imbricate, strongly ventricose at base, antheridia 1--3 per bract. Gynoecia on terminal or separate
branches, bracts in 2--5 pairs, similar but larger than stem leaves; perianth ± flattened,
ovoid to obovoid, short, contracted to mouth, 6--9 plicate, upper 1/3--1/4
narrowed, mouth denticulate with 1--rarely 2---celled rigid teeth. Sporophyte not seen.
Rock
outcrops; moderate to high elevations; Nfld. and Labr. (Nfld.),
Ont.; N.C., Minn., Tenn.; Central America (Costa Rica); Asia (Japan).
5. Diplophyllum obtusifolium (Hooker) Dumortier, Recueil d'Observations sur les
Jungermanniacées,
16. 1835
Jungermannia obtusifolia Hooker,
Brit. Jungerm., plate 26, 1816
Plants in patches or among other
bryophytes, light green in shade to yellow, chestnut or red-brown in sun, in
direct sunlight, locally vinaceous pigmented at
ventral lobe bases. Stems 0.8--1 cm, branches abundant,
prostrate; leafy shoots 1.5--2.5
mm wide; cortex 1--2 layers thick, rhizoids abundant
on ventral surface of stem, abundant, often near ventral leaf bases. Leaves contiguous to imbricate,
often dorsally secund, lingulate, often truncate at the apex, but
occasionally ± apiculate; ventral lobes 0.5--1.5 x 0.35--0.5 mm, ca. 2 x as large
as dorsal lobes, lingulate, nearly or quite parallel-sided, often weakly
falcate, diverging 75--90º from stem, proximal margins distinctly irregularly
denticulate at proximal base, ± finely denticulate at apex, apices
rounded-truncate; dorsal lobes 0.3--0.9 x 0.4--0.7 mm, ca. 0.6 x the
ventral lobe in area, diverging from the stem at ± 30º and often directed
almost to the stem apex, ± appressed to stem and lobe, lingulate to
ovate-lingulate, margins finely denticulate, at least near apex; cells firm,
rather opaque, ca. 10 \um at leaf tip,
marginal cells 8--12 \um, medial and submedial
cells sublinear, 2--4:1, ca. 40--64 x 12--16 \um;
cuticle usually ± papillose; oil bodies spheric
to ovoid, 2--5 per medial cell. Specialized asexual reproduction usually
absent or very rarely by stellate, 1-celled gemmae. Sexual condition paroicous. Androecia bracts in 3--5 pairs,
similar to stem leaves, lingulate lobes, antheridia 1--3 per bract. Gynoecia bracts in 3--5 pairs, similar
stem leaves; perianth plicate, short-clavate to
oblong clavate, contracted to mouth, shallowly 5--6-lobulate, shallow lobes
bearing 1--3, mostly 1-celled teeth; Sporophyte
capsule ovoid. Spores 8--12
\um, delicately papillose-verrucose.
Moist
shaded soil banks and boulders; 15--600 m; Greenland; B.C.; Alaska, Calif.,
Oreg., Wash.; Europe; Asia (Japan).
Diplophyllum
obtusifolium is distributed across temperate and subarctic
regions.
6. Diplophyllum andrewsii
A. Evans, Bryologist 25: 28, plate 1,
figs. 1--11, 1922
Plants in small patches, ± shining
pale to yellow-green, translucent in shade, becoming clear, translucent,
often golden brown in sun. Stems prostrate with apices ascending,
0.3--0.8 cm,
0.15--0.22 mm
in diameter, irregularly and sparingly branched below perianth,
cortex 1--2 layers thick; leafy shoots 1.3--2.4 mm wide; rhizoids
dense on prostrate stems. Leaves
with entire margins, loosely to closely imbricate in the shade, when
moist, nearly flat and spreading horizontally, strongly dorsally secund and connivent in the sun, the opposed ventral lobes with
their tips often closely overlapping; dorsal lobes when moist suberect to
obliquely spreading 10--45º angle with the stem, when dry oblong--ovate to
short-lingulate, squarrose, broadly rounded at apex, 0.65--0.8 x 0.4--0.45 mm, up to 0.5 x as
large as ventral lobe; ventral lobes widely spreading, at a 80--110º angle
with the stem, oblong-lingulate, 1--2 x 0.5--0.6 mm, 2:1, ± strongly
concave, margins raised at least in sun forms, keel short suberect, nearly
straight to slightly arching outward near apex; marginal cells in several
rows, isodiametric, thick-walled, ca. 10 \um; medial cells thin-walled with
indistinct concave-sided trigones, 20--23 x 14--16 \um; cuticle almost
smooth, papillae of medial cells very pale, minute and inconspicuous; oil
bodies, homogenous, glistening, spheric to
short-ovoid in distal fourth of lobes and
submarginal cells, 4--6 per cell, 6--10 in medial and submedial cells. Specialized
asexual reproduction by fragmented branches, gemmae very rare, small, spheric to subspheric, ±
brownish. Sexual condition
autoicous, abundantly fertile. Androecia
bracts in 4--6 pairs, similar to stem leaves, antheridia 1 per bract. Gynoecia on short lateral branches,
bracts in 2--5 pairs of increasingly larger leaves, similar to but somewhat
larger than stem leaves; perianth at maturity 1/2
exserted, oblong to fusiform, irregularly 8--10-plicate, barely dorsiventrally flattened, gradually contracted to
somewhat pointed, mouth irregularly and shallowly 10 or more lobed, lobes
acuminate, ending in cilia formed by 2--5 elongated cells, sides of lobes
with 1--2-celled sharp teeth. Sporophyte
unknown in North America.
Shaded
loamy soil of eroding banks along streams, rock ledges and crevices; 270--1700 m; N.C., S.C., Tenn., Va.; Japan.
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