BFNA Title: Diplophyllum
 Author: P. L. Redfearn, Jr. 
Date: July 23, 2008
Edit Level: R 
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

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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.