BFNA Title: Solenostoma
XX. SOLENOSTOMA Mitten, emend. Zerov, J. Linn. Soc. Bot., 8: 51 1865 * [From Greek solen, canal and stoma, mouth, alluding to perianth suddenly narrowed to mouth]
Jungermannia subg. Solenostoma (Mitten) Amakawa
Plants 3--20 mm x 0.5--3.4 mm, ascending to erect, rarely creeping, greenish, pale green yellowish, yellowish brownish to dirty green, in the most species with characteristically red or purple coloration (at least as a tint at the tips of perianthia and leaf rim). Stem orbicular to transversely oval in cross section 100--520 \um in width and 80--500 \um in height, dorsal surface cells mostly subquadrate, more rarely shortly elongate with walls colorless to slightly brownish, trigones mostly small and concave, cuticle smooth to striolate; in the stem cross section outer cells smaller and with thicker cells. Rhizoids mostly dense, rarely scattered, in bundles, colorless to brownish, rarely reddish or purplish. Leaves contiguous to imbricate, subtransversely to (more rarely) obliquely inserted, dorsally decurrent or not, ventrally insertion line arcuate, not or long-decurrent; cells at midleaf mostly thin-walled, walls colorless to yellowish, never deep brownish or orange-brown, trigones mostly distinct, concave to convex. Sexual condition paroicous or dioicous. Specialized asexual reproduction present in one species only, by endogenous 1--2-celled oval gemmae. Perianth terminal on main axis, innovations rare, emergent from bracts for 1/4--3/4 of its length, tubular obtrapezoidal, 1-stratose to base or sometimes 2-stratose in proximal 1/5(--1/3) of the length, mostly suddenly contracted to the beaked mouth and distinctly 3--5-keeled (1--2- dorsal, 0--2- lateral, 1--2- ventral), perianth mouth crenulate, composed by elongate cells, cells in mid perianth mostly isodiametric, hexagonal to rectangular, never linear, trigones mostly distinct and convex; perigynium absent, or, if present, short, to 1/3 of perianth length. Androecia intercalary, in (1--)2--5 pairs of bracts, with 1--3(--4) antheridia per bract, stalk 2-seriate. Capsule spheric to shortly ellipsoidal, 2-stratose. Elaters 2-spiral, ca. 100 x 7.5 \um, 2-spiral, with or without homogenous ends. Spores brown to brown reddish, spheric, finely papillose, 12--19 \um.
ca. 70 (7 in the flora): North America, Central America, South America,
Crum, H. A. 1991. Liverworts and
Hornworts of southern
1. Plants paroicous, leaves without distinct rim of swollen cells along margin, midleaf cells mostly with trigones moderate in size.
2. Plants mostly erect, more rarely ascending, green in shady places, to mostly sepia-brown in color, never purplish, rhizoids originated from ventral side of stem only … 7. Solenostoma sphaerocarpum
2. Plants ascending to creeping, green in shady places, to brown-red to purplish in color, rhizoids originated from both ventral side of stem and from some cells in lower half of leaves … 2. Solenostoma confertissimum
1. Plants dioicous, leaves with or without distinct rim of swollen cells along margin, midleaf cells commonly with trigones moderate to large in size and bulging, or trigones indistinct.
3. Plants whitish to yellowish, without traces of red pigmentation, with the exception of perianth beaks, plants almost constantly with endogenous gemmae present within unfertilized perianths, oil-bodies brown to brownish, 1(--2) per midleaf cell …1. Solenostoma caespicitium
3. Plants from yellowish to deep brown-red, gemmae absent, oil-bodies grayish to nearly colorless, more than 3 per midleaf cell.
4. Plants lax,
large (mostly more than
4. Plants not lax, small (mostly less than 2 mm wide), leaf margin not undulate, leaves horizontally spreading, not sheathing the stem or sheathing only near the leaf base, not blackish, commonly reddish to red brown, the stem not whitish, plants creeping to loosely ascending.
5. Leaves bordered with swollen cells with equally thickened cell walls, size of marginal cells at least two times larger than midleaf cells; rarely leaves not bordered, but in the latter case trigones in the leaf are not developed … Solenostoma gracillimum
5. Leaves not bordered with a rim of swollen subequally thickened cell walls, size of marginal cells nearly the same or only slightly larger than midleaf cells.
6. Leaves bordered with rim of strongly unequally thick-walled cells, with large, sometimes semi-confluent trigones, size of marginal and midleaf cells nearly the same … 6. Solenostoma rubrum
6. Leaves not bordered, cell walls evenly thin, with moderate to large convex trigones … 5. Solenostoma appalachianum
1. Solenostoma caespiticium (Lindenberg) Stephani, Sp. Hep., 2: 57
Jungermannia caespiticia Lindenberg Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 14(Suppl.): 67. 1829
On soil near stream, field banks, clayey roadsides; 0--500 m; B.C.; Alaska, N.Y.
2. Solenostoma confertissimum (Nees) Schljakov, Pechen. Mkhi Severa SSSR 4: 5.1 1981
Jungermannia confertissima Nees, Naturgesch. Eur. Leberm. 1: 277, 291. 1833; J. pyriflora subsp. purpurea (R.M. Schuster & Damsholt) Stotler & Crandall-Stotler; J. pyriflora subsp. purpurea var. innovata (R.M. Schuster & Damsholt) Stotler & Crandal-Stotler; Solenostoma levieri (Stephani) Stephani; S. pyriflorum var. innovatum R. M. Schuster & Damsholt; S. pyriflorum subsp. purpureum R. M. Schuster & Damsholt
On moist soil and
wet rocks near streams frequently associated with mosses and other hepatics;
0--1100 m or higher; Greenland; B.C., Que. ; Alaska, Calif.,
3. Solenostoma fusiforme (Stephani) R. M. Schuster, Hepat. Anth. N. Amer. 2: 944. 1969
Nardia fusiformis Stephani, Bull. Herb. Boiss., 5(2): 99. 1897; Jungermannia fusiformis (Stephani) Stephani; J. koreana (Stephani) Amakawa; Plectocolea koreana (Stephani) S. Hattori; Solenostoma koreanum Stephani
Plants 15--25 x 1--2.8 mm, ascending to erect in dense patches, pale green to whitish, with characteristically brownish to reddish brown colored leaf margins. Stem 315--420 x 280--390 \um, oval in cross section, dorsal surface cells thin- to slightly thick-walled, cells obliquely rectangular to oblong-hexagonal, with small concave or indistinct trigones, 100--125 x 17--33 \um. Rhizoids dense, colorless to brownish, in indistinct fascicles at 90º with stem; in the stem cross section outer layer 2-stratose mostly composed by moderately thick-walled cells, ca. 12--25 \um in diameter, inner cells thin-walled with small to indistinct trigones, in central part nearly the same size with outer layer, but between center and outer layer larger up to 45 \um in diameter. Leaves distant to contiguous, inserted at angle of 10--40° with axis, dorsally decurrent for ca. stem width, ventrally subtransversely inserted, not or barely decurrent, (700--)900--1750 x (665--)1120--2135 \um (1:0.6--0.95(--1.2)), transversely oval to rounded- widely triangular, mostly flattened to slightly concave, moderately and large-sized frequently undulate at margin; mostly with distinct brown to reddish brown and purplish colored rim along margin; cells in the midleaf thin-walled or slightly thickened, colorless, subisodiametric to oblong hexagonal, 30--45 x 25--38 \um, trigones small and concave; near margin 28--33 \um, with mostly thickened, but not strongly, or (rarely) thin-walled but with thickened external wall, brownish to reddish brown or, occasionally, colorless; in basal part cells oblong rectangular to oblong hexagonal, 50--100 x 30--43 \um; cuticle smooth or very loosely striolate. Sexual condition dioicous. Sporophyte unknown.
On wet rocks and
soil near sluggishly flowing streams, more rarely in bogs; 0--500 m; B.C.,
4. Solenostoma gracillimum (Smith) R. M. Schuster, Hp. Anthocerotae N. Amer. 2: 972. 1969
Jungermannia gracillima Smith, Engl. Bot. 32: tab. 2238. 1811; J. crenulata Smith non J. crenulata Schmidel; Solenostoma crenulatum (Smith) Mitten
Plants 3--10 x 0.4--0.9 (fertile up to 1 mm) mm, prostrate to (fertile) ascending, yellowish brownish to pale green to pale yellowish and brown; female plants yellow red to brownish purplish. Stem (100--)175--245 \um in width and (90--)150--220 \um in height, branching lateral as normal branch or slender innovations; dorsal surface cells 35--75 x 17--25 \um, rectangular to oblong hexagonal and obliquely rectangular, walls thin; cells in the stem cross section nearly the same size in outer and inner layers, but outer with cell walls thickened with yellow to brownish colored walls, inner cells more or less thin-walled, with walls colorless, ca. 15--20 \um in diameter. Rhizoids sparse, colorless, in indistinct fascicles, tawny or decurrent down the stem. Leaves distant to subimbricate (especially in area 5--7 pairs below the perianth), inserted at angle (10--)15--25º, although sometimes to subtransverse in leaves proximal to female bracts; (190--)470--900 x (200--)500--940 \um (1: (0.7--)0.95), orbicular to rounded-triangular, lingulate and reniform, widest at proximal 1/3 of leaf length, more rarely near the middle; cells in the midleaf (22--)30--45 x (17--)25--28 \um, thin-walled, subisodiametric, mostly 5--6-gonal, trigones indistinct to small; along margin (25--)30--70 \um, very thick-walled or not differs from midleaf cells (then leptodermous, without trigones to slightly equally thickened), walls brownish, cuticle striolate at proximal 1/2 of the leaf. Sexual condition dioicous. Perianth terminal on main axis, no innovations, emergent for 1/3--1/2 of its length, conical, tubular-conical, ellipsoidal to short-clavate, with the suddenly tightened mouth 4--5-plicae (1--2- dorsal, 2- lateral, 1- ventral), perianth mouth crenulate, composed elongate thick-walled cells, slightly dilated on the external end, perianth in the middle part rectangular to isodiametric, ca. 30--45 x 18--25 \um, cuticle distinctly striolate in lower half; perigynium ca. 1/4 of the perianth length or smaller; bracts not differentiable from sterile leaves, slightly larger only, in 1 pair, ca. 950 x 950--1070 \um. Androecia intercalary, spicate, in 2--3 pairs of bracts, strongly inflated in the base, with incurved margin of bracts, with 1 antheridium per bract, body brownish, shortly ellipsoidal ca. 125 x 112 \um, stalk 2-seriate, ca. 33 x 17 \um. Seta ca. 3000--5000 x 125 \um. Capsule shortly ellipsoidal, 2-stratose, inner cells long rectangular 38--63 x 7--13 \um, with 8--10 annular thickenings, outer cells subquadrate, ca. 25--30 x 17--35 \um with 1--2 nodular thickenings in both vertical and horizontal walls, elaters 2-spiral, ca. 100 x 7.5 \um, with homogenous endings ca. 10 \um length, spores red-brown, spheric, finely papillose, 10--11 \um in diameter. Elaters 600--800 x 6-7 \um. Spores 10--12 \um, red brown, papillose.
fine-grained soil along streams and (more frequently) in human disturbed areas;
0--500; N.B., Nfld. and Labr. (Nfld.), N.S., Ont., P.E.I., Que.;
There are two forms of Solenostoma gracillimum in the flora area. The fo. crenulatum (Mitten) Bakalin is distinguished from the widespread fo. gracillimum by leaves with the rim of swollen thick-walled cells, 1.5--2 times larger than midleaf cells and is known only from Arkansas, Kentucky, Texas and distribution in Eurasia.
5. Solenostoma appalachianum R. M. Schuster ex Bakalin Arctoa, 23: 127. 2014
In moist rock
crevices or on rocky surfaces, rarely on bare soil in human disturbed areas;
700--1000 m; Ga.,
Solenostoma appalachianum has been misidentified in the past under the names Solenostoma pyriflorum or Jungermannia pyriflora.
6. Solenostoma rubrum (Gottsche ex Underwood) R. M. Schuster, Hepat. Anthocerotae N. Amer. 2: 975. 1969 E
Jungermannia rubra Gottsche ex Underwood, Bot. Gaz. 13: 113. 1888
Plants 3--8 x 0.7--1.8 mm, prostrate to (mainly) ascending and erect, green, green purplish (due to purple-colored ventral base of leaves) to yellowish pale green, yellowish brown, brownish, brownish reddish, with markedly purple colored perianth beak and tops of ascending erect shoots; female plants the same size with sterile ones or slightly larger, male are wider due to incurved apices of the bracts. Stem 210--370 \um in width and 160--300 \um in height; dorsal surface cells rectangular, ca. 74--100 x 25--38 \um thin-walled trigones indistinct, cuticle striolate-papillose; in the stem cross section outer cells smaller than inner ones, ca. 17--25 \um in diameter, walls slightly thickened (external wall evidently thickened), yellowish, trigones distinct, concave, inner cells 5-gonal to polygonal, ca. 33--50 \um in diameter, thin-walled, walls colorless, trigones small to indistinct. Rhizoids sparse to common, mostly confined to area near ventral base of leaves and commonly absent in distal 3--4 pairs of leaves, colorless to pale grayish and brownish, in indistinct fascicles decurrent down the stem. Leaves distant to contiguous and subimbricate, inserted at angle ca. 45--65º with axis, dorsally decurrent for 1/4 of the stem width, ventrally inserted subtransversely, insertion line arcuate, barely decurrent (in some shoots are inserted at angle ca. 80--85° with the axis and not decurrent both dorsally and ventrally); 630--1050 x 700--1225 \um (1:0.6--1), suborbicular, transversely oval to orbicular-lingulate and orbicular-ovate, widest near the middle, concave to channeled, sometimes deflexed away the stem; cells in the midleaf thin-walled, subisodiametric, mostly 5--6-gonal, 30--45 x 30--60 \um, walls colorless to purplish, trigones distinct, concave to triangle; along margin 20--45 \um, with external wall only clearly thickened (rarely walls equally thickened), sometimes, with visible median lamina in marginal cell walls, trigones distinct, triangle, cuticle smooth. Sexual condition dioicous. Perianth terminal on main axis, no innovations, emergent for 2/3 of its length, 1470--1750 x 985--1050 \um, clavate to shortly fusiform, with 4 distinct plicae (1-dorsal, 2-lateral, 1-ventral), suddenly tightened to the beaked mouth, perianth mouth crenulate, perianth in distal part 1-stratose, cells subisodiametric to oblong, 25--33 x 38--50 \um, trigones distinct to convex, in the middle part subisodiametric, ca. 33--50 x 30--38 \um, walls thin, trigones distinct, triangle to convex, cuticle smooth, in basal part 2-stratose, cells rectangular to oblique rectangular, 100--150 x 28--38 \um, trigones small and concave, cuticle striolate-papillose; perigynium small, lesser than 1/6 of the perianth length; bracts just below perianth, in 1 pair, ca. 720--900 x 950--100 \um, lingulate to obtrapezoidal, frequently shortly 2-lobate, clinging to perianth in its basal part, but deflexed distally. Androecia intercalary, spicate due to deflexed tops of bracts, in 3--5 pairs of bracts, ca. 630 x 875 \um, orbicular to transversely oval, strongly inflated at the base and cupped, but with incurved margin, with 2 antheridia per bract, antheridial body shortly elliptical to spheric, ca. 110--140 x 95--140 \um or smaller (probably immature), stalk 2-seriate. Seta ca. 3000 \um x 200 \um. Capsule brown, shortly ellipsoidal, ca. 500--625 x 490--500 \um; inner cells 45--60 x 10--15 \um, oblong-rectangular to irregular in shape, with 7--8 annular thickenings; outer cells rectangular, 25--50 x 15--23 \um with 3--4 nodular thickenings in each of vertical wall and 0--1 on horizontal wall. Elaters reddish brown, 2-spiral ca. 112 x 7.5 \um, homogenous narrowed endings from each side ca. 13--15 \um in the length. Spores brown to reddish brown, papillose, ca. 15--16 \um.
On moist soil
along stream, clayey roadsides; 0--2160 m; Alta., B.C.;
7. Solenostoma sphaerocarpum (Hooker) Stephani, Sp. Hepat. 2: 61. 1901
Jungermannia sphaerocarpa Hooker, Brit. Jungermann., pl. 74. 1815
Plants 3--7 x 0.35--2 mm, erect to ascending, sepia to brownish yellow and grayish brown, without traces of red pigmentation. Stem ca. 175--300 \um in diameter, branching not seen; dorsal surface cells 33--45 x 22--30 \um, subrectangular, cuticle obscurely striolate; in the stem cross section cells ca. 20--25 \um in diameter, walls brownish, mostly thickened, trigones small but distinct, concave. Rhizoids numerous, dirty grayish to almost colorless, in indistinct fascicles, mostly erect, more rarely decurrent. Leaves contiguous to distant, subtransversely inserted, 400--700 x 550--900 \um (1:0.6--0.8), widest in middle or just below, concave to almost flat; cells in midleaf 20--25 x 19--25 \um, lumen rounded to hexagonal, walls thin to obscurely thickened, brownish, trigones large and convex; cuticle striolate in lower 1/3. Oil-bodies 5--6 per cell, finely granulate, brownish-grayish, oval in projection, 5--11 x 4--7 \um. Sexual condition paroicous. Perianth terminal on main axis, innovations lateral, 1--2 per gynoecium, rarely present; emergent for 1/4--2/3 of its length, 1540--1750 x 1540--1540 \um, tubular with the suddenly tightened beaked mouth, with 3--4 distinct plicae (1--2-dorsal, 2-ventral), perianth mouth crenulate, perianth mainly 1-stratose, but 2-stratose in basal portion, in the middle part of perianth cells 28--35 x 17--23 \um, subisodiametric to oblong, mostly hexagonal; perigynium up to 1/3 of the perianth length; bracts almost not differentiated from sterile leaves, only slightly larger, in 1 pair, ca. 750--800 x 1000--1250 \um, concave to flattened, rarely obscurely undulate at margin or slightly retuse. Androecia below perichaetium, in 1--3 pair of bracts, bracts similar to sterile leaves, slightly inflated near the base, with 2--3 antheridia per bract, stalk 2-seriate, ca. 43 x 17 \um. Seta ca. 1500--2000 x 120 \um. Capsule oval ca. 600--700 x 400 \um, inner cells 50--70 x 12--15 \um, frequently with flexuous walls, with 6--8 annular thickenings, outer cells nearly subquadrate, ca. 30 x 28 \um with 2 nodular thickenings in vertical wall. Elaters 2-spiral, ca. 150 x 8 \um. Spores finely papillose, ca. 17 \um.
On moist soil and
rocks near streams, on wet fine-grained soil in anthropogenic or natively
disturbed areas; 0--3100m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.),