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BFNA Title: Solenostoma |
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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] Vadim Bakalin 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. Species
ca. 70 (7 in the flora): North America, Central America, South America,
Eurasia, Africa, SELECTED
REFERENCES 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 Plants 0.5-- 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 Plants 3-- 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.,
Colo., 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. On
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 Plants 3-- 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.),
N.W.T., |
