BFNA Title: Plectocolea |
XX. PLECTOCOLEA
(Mitten) Mitten, Fl. Vit., 405. 1871 * [Greek, pleko, turbinate,
and koleos, vagina, alluding to the turbinate
distal perianth] Solenostoma subg. Plectocolea Mitten, J.
Proc. Linn. Soc. Bot. 8: 156 1865; Jungermannia Linnaeus subg. Plectocolea (Mitten)
Amakawa Plants 3--150 x 0.6--5
mm, prostrate to ascending or erect, yellowish brown, brownish, reddish
brown, or deep green . Stem transversely elliptic in cross section, branching lateral
intercalary or ventral, rarely as postical innovations and scale-like leaved strongly
rhizogenous branches; dorsal surface cells
rectangular to usually linear, thin- to thick-walled with distinct or
indistinct trigones, cuticle mostly striolate; stem cross section mostly
differentiated in 2--3 layers, with the outer as a hyaloderm,
or if hyaloderm absent a scleroderm,
in some species the differentiation may be unclear to virtually absent. Rhizoids red, red-brown to purple,
blackish brown to green, almost always brown with purple or purplish
pigmentation at least as traces. Leaves
distant to contiguous, never imbricate, obliquely inserted, mostly distinctly
decurrent dorsally and sometimes ventrally, midleaf cells subisodiametric,
mostly 5--6-gonal, walls colorless to yellowish and brownish, slightly
thickened to evidently thickened or thin-walled, trigones distinct in most
species, mostly concave, rarely large and convex. Sexual condition
dioicous or paroicous. Perianth
terminal on main axis, conical to tubular and pyriform, 2-stratose in
proximal third, of long-rectangular to semi-linear cells at mid perianth. Perigynium always well-developed reaching 1/2--2/3 times the perianth length. Androecia intercalary, with 1--10
pairs of bracts (frequently androecial branch with dead androecia distally),
1--3 antheridia per bract, stalk 2-seriate. Capsule wall 2-stratose. Elaters
2-spiralled, always with homogenous ends, more or less elongate, 1/5--1/10
times the elater length. Species
ca. 50 ( The
exact number of Plectocolea species is unknown because many taxa
not yet revised have been described from SELECTED
REFERENCES Doyle W. T. and R. E. Stotler. 2006.
Contribution toward a bryoflora of California III. Keys and Annotated species
Catalogue for Liverworts and Hornworts. Madroño.53: 89--197. Damsholt, K. and J. Váňa.
1977. The genus Jungermannia L. emend. Dumort. (Hepaticae) in 1. Plants
dioicous, leaves with distinct rim of swollen cells, far eastern portion of
flora … 1. Plectocolea crenuliformis 1. Plants paroicous
or dioicous, leaves without distinct rim of swollen cells rim, although
marginal cells may be more intensively colored. 2. Plants
paroicous. 3. Plants pellucid, merely soft, light green to yellowish
green, leaves obliquely oriented, leaf margin (especially in distal part of plant)
frequently undulate. Perigynium low, ca. 1/3 times the perianth length or
less; rhizoids purplish … 2. Plectocolea fossombronioides 3. Plants not pellucid, merely rigid, deep green to
brownish green and (when small) almost constantly with marked traces of
purple coloration; leaves not undulate, more or less vertically oriented at
least in normally developed shoots. Perigynium more than 1/2 times perianth
length; rhizoids purple to brownish. 4. Plants large, wider than 2.2 mm, rhizoids purple, leaves
vertically oriented … 4. Plectocolea obovata 4. Plants small, less than 2 mm in width, rhizoids purplish
to brownish, leaves vertically to obliquely oriented
… 7. Plectocolea subelliptica 2. Plants dioicous. 5. Plants pale
green to pale yellowish brownish to brownish-green, leaves in larger plants
frequently undulate at margin, shoots mostly less than 5. Plants deep to
dark green, brown green, purplish brown and blackish green, leaves not or
slightly undulate at margin (in plants larger than 6. Plants 1.4-- 6. Plants 2.1--4.8 mm wide, perigynium 2 times longer than
perianth, oil-bodies in the midleaf commonly botryoidal
… 6. Plectocolea schusteriana
Jungermannia crenuliformis Austin, Bull. Torrey Bot. Club 3: 10 1872; Solenostoma crenuliforme (Austin)
Stephani Plants 3--15(--20) x 1.1--2.5
mm, prostrate to ascending, yellowish brown, brownish to brown reddish, with
characteristically more deeply colored leaf margin, rarely in shady places pale
greenish to whitish and very soft and gentle; female plants larger than the sterile.
Stem 105--360(--385) \um in width
and 80--250(--305) \um in height, branching lateral intercalary or ventral,
the latter rare; dorsal surface cells 63--175 x (12--)15--25 \um, linear,
rectangular to obliquely rectangular, walls thin to very thick and brownish
(ca. 2.5--3.5 \um in thickness), trigones indistinct, cuticle loosely
striolate; in the stem cross section there are at least four variants of
structure: (1) three-layered with outer cells (hyaloderm)
ca. 17--30 \um in diameter, thin-walled, walls colorless, middle stratum (scleroderm), cells ca. 12--15 \um in diameter,
thick-walled, walls yellowish, inner cells ca. 12--23 \um in diameter,
slightly thick-walled to thin-walled; (2) in the stem cross section there are
2 strata: outer with slightly thick-walled cells, ca. 25--27 \um in diameter,
inner with cell wall thin to slightly thickened, cells mostly hexagonal, ca.
25--37 \um in diameter, cell walls colorless; (3) no distinct differentiation
in the stem cross section, walls become thicker to dorsal side, outer cells
larger, ca. 25-- On circum-neutral to acidic rocks, sandstone, along shaded
rocky streams, or in disturbed areas in forested lowlands; 0--550 m; Ont.,
Que.; 2. Plectocolea fossombronioides (Austin) Mitten,
Trans. Linn. Soc. London, Bot., ser 2. 3: 198 1891 Jungermannia fossombronioides Austin, Proc. Acad. Nat. Sci. Philadelphia 1869: 220. 1869;
Solenostoma fossombronioides
(Austin) R. M. Schuster Plants very soft, 3--6 x
0.7--1 mm, prostrate to ascending (near apex of fertile shoot), pale
brownish, whitish, yellowish brown; fertile plants slightly larger than
sterile. Stem 105--175 \um in
width and 100--165 \um in height, dorsal surface cells ca. 70--87 x 25--30 \um,
thin-walled or slightly thickened, trigones indistinct; stem cross section
composed of mostly thin-walled cells, mostly penta-
to hexagonal in cross section, ca. 20--38 \um in diameter, smaller near
ventral side, walls colorless to brownish, trigones absent or barely visible.
Rhizoids dense, brown-red to (more rare) purplish and pale brownish, rigid, separated
one from other, or sometimes in indistinct fascicles, originated at 90º with
the stem. Leaves distant to
contiguous, inserted at angle 15--20º with stem, dorsally shortly decurrent
(not more than 1/5 of stem width), ventrally inserted at angle 50--60º with
axis, not decurrent; 600--700 x 490--600 \um (0.8--0.9:1), lingulate to
ovate, widest between 1/2 and 1/3 of leaf length, flattened to obscurely
concave, the biggest leaves frequently undulate at margin; cells at mid leaf
thin-walled, 38--60 x 35--43 \um, walls brownish to colorless, trigones
small, concave; along margin 25--35 \um, slightly elongated along margin,
walls thin, colorless; cuticle smooth everywhere. Oil-bodies 2--5(--8) per cell, 6--8 x 8--22 \um. Sexual
condition paroicous, but occasionally with some branches androecious only (heteroicous).
Perianth terminal on main axis, no
innovations, emergent for 1/3--2/3 of its length, tubular, pentaplicatae when mature (1-dorsal, 2-lateral,
2-ventral) or pluriplicate, obscurely tightened to the mouth, perianth
composed by uniformly elongated cells, ca. 70--85 x 25--33 \um, walls thin,
trigones small to indistinct, cuticle smooth. Perigynium 3/4--1/1 of the perianth length, with 2 pairs of
bracts; bracts just below perianth, in 2 pair, the same shape as male bracts.
Androecia just below female bracts
(or, occasionally only in separate branches, in heteroicous
plants), in 2--3(--4) pairs of bracts (if in separate branches then 3--4
pairs of bracts), with 1--2 antheridia per bract, antheridial
body shortly elliptical to spherical, ca. 120--170 x 120--170 \um, stalk 2-seriate,
ca. 38 x 25 \um; bracts strongly inflated near the base with margin
(especially the pair nearest to the perichaetium) deflexed away of the
perianth. Seta ca. 150 \um in
diameter. Capsule shortly
ellipsoidal, ca. 700 x 630 \um, 2-stratose, inner cells, long-rectangular
50--100 x 7.5--13 \um, with 5--8 annular thickenings, outer cells subquadrate
to rectangular, ca. 33--55 x 40--43 \um with 2--3 nodular thickenings in
vertical wall, and 0--2 on the horizontal one. Elaters 2-spiral, ca. 165 x 7.5 \um, with homogenous endings, ca.
30 \um length, spores brown to brown reddish, spherical, finely papillose,
15--16 \um in diameter. Soil-covered
rocks along streams, occasionally pioneer on sedimentary or slightly
metamorphosed rocks, temperate species distributed in forested areas; 0--900;
Conn., D.C., Ga., Ill., Ind., Kans., Minn., Miss., N.J., N.C., Ohio, S.C.,
W.Va. The capsule of Plectocolea fossombronioides
matures in mid April. 3. Plectocolea hyalina (Lyell) Mitten,
Trans. Linn. Soc. London, Bot. Ser. 2, 3: 198. 1891 Jungermannia hyalina Lyell, Brit.
Jungermann. Pl. 63. 1814; Solenostoma hyalinum (Lyell) Mitten Plants 3--8 x 0.7-- On wet soil or
soil covered rocks, sometimes among mosses in wet mossy tundras;
0--2500; 4. Plectocolea obovata
(Nees) Mitten
Fl. Vit., 405 1873 Jungermannia obovata Nees Naturgesch.
Eur. Leberm., 1: 332 1833 Solenostoma obovatum (Nees) R.M. Schuster Hep. Anth.
N. Amer., 2: 1007 1969 Plants 4-- On shaded wet
rocks and soil near streams, rarely submerged, mostly in tundra zone or belt;
0--2000 m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), Que.; 5. Plectocolea obscura (A. Evans) A.
Evans in Buch, A. Evans & Verdoorn, Ann. Bryol.
10: 42. 1938 Nardia obscura
A. Evans, Rhodora 21: 159. 1919; Jungermannia evansii Váňa Plants 4--8 x 1.4--2.1
mm, prostrate to ascending, green brownish, with light purplish traces near
apices; female plants larger. Stem
200--400 \um in width and 120--300 \um in height, dorsal surface cells long-rectangular
to obliquely rectangular, ca. 75--150 x 15--23 \um, mostly thin-walled with
indistinct trigones; in the stem cross section there are 3 strata: outer (hyaloderm) with thin external wall and moderately
thickened other walls, ca. 12--25 \um in diameter, medium (scleroderm) composed by thick-walled cells, walls
yellowish in color, ca. 12--20 \um in diameter, inner with cell wall thin to
moderately thickened, cells mostly polygonal, ca. 12--22 \um in diameter,
cell walls colorless. Rhizoids purple,
rigid, in a bundle or separated one from other, angled at 90º with stem. Leaves distant to contiguous,
inserted at angle of 15--30° with axis, dorsally shortly, but distinctly
decurrent; 1050--1225 x 980--1190 \um (0.85--0.95:1), oval to ovate, widest
at midleaf or slightly below, with rounded to retuse apex, adherent to the
stem in the base and flattened and erect above; cells at mid leaf
thin-walled, subisodiametric 28--38 x 25--33 \um, trigones convex; along
margin 15--23 \um, thin-walled with convex trigones; cuticle distinctly
papillose in the proximal third of the leaf, in distal parts papillae very
obscure or absent. Oil-bodies 2--5(--10)
per cell, ovoid to ellipsoidal, 5--9 x 6--14 \um. Sexual condition dioicous.
Perianth terminal on main axis,
immersed, conical loosely pluriplicate with 3--4 main plicae
those are undulate to the mouth, ca. 700 x 600 \um, perianth mouth crenulate,
perianth in distal part 1-stratose, cells mostly rectangular, 25--43 x 17--18
\um, walls thin, trigones small, cuticle smooth or finely papillose, perianth
in proximal part 2-stratose, cells mostly oblong rectangular, 38--50 x 10--25
\um, walls thin to thickened, trigones small and concave, cuticle distinctly
striolate papillose. Perigynium ca.
1/1--3/2 of the perianth length, with 1--1.5 pairs of bracts; bracts just
below perianth, in 1--2 pairs, lingulate, adherent to perianth and perigynium
at the base and deflexed away the perianth in distal half, 1470--1750 x
1015--1470 (0.7--0.9:1). Androecia
intercalary in 2--3 pairs of bracts, with 1--3 antheridia per bract, stalk 2-seriate,
ca. 50--63 x 15--18 \um; bracts strongly inflated in base, but deflexed away
the stem in distal part. Sporophytes unknown. On wet acidic
rocks near streams, largely a montane in subalpine habitats; 0--2000 m;
Greenland; B.C.; Alaska, Ga., 6. Plectocolea schusteriana (J. D. Godfrey
& G. Godfrey) Bakalin, Bot. Pacif. 3: 92. 2014 E Jungermannia schusterana J. D. Godfrey & G. Godfrey, J. Hattori Bot. Lab. 46:
109, fig. 1--3. 1979; Solenostoma schusterianum
(J. D. Godfrey & G. Godfrey) Váňa, Hentschel & J. Heinrichs Plants 20--150 x 2.1--5
mm, ascending to erect, deep green, brownish deep green and green brown and green-purple
(due to purple colored ventral bases of leaves); female plants slightly
larger than male and sterile and have mainly retuse leaves (even far down of
perianth). Stem 280--525 \um in
width and 220--400 \um in height, dorsal surface cells thin-walled, mostly
elongate-rectangular to obliquely rectangular and linear, 70--125 x 15--20
\um, trigones small and concave, cuticle distinctly striolate; in the stem
cross section there are 3 strata: outer (hyaloderm)
moderately thick-walled to almost thin-walled, ca. 12--25 \um in diameter,
walls brown (external wall more deeply) to purplish and purple colored,
medium (scleroderm) very thick-walled with commonly
visible median lamina of cell wall, ca. 17--25 \um in diameter, inner
thin-walled to moderately thick-walled, ca. 17--38 \um in diameter, with
indistinct trigones and walls mostly colorless. Rhizoids uncommon in male and sterile shoots, but more or more dense
on female, purple to brown-purple, in the bundle or separated one from other
or in indistinct fascicles. Leaves
distant to contiguous, inserted at angle of 50° with axis, dorsally decurrent
for 1/2--2/2 of the stem width, ventrally subtransversely inserted, line
insertion arcuate, decurrent for 1/2--2/2 of stem width; (875--)1100--2400 x
(830--)1000--2000 \um (0.6--0.9(--15):1), oval to ovate-oval, obliquely
ovate, widest slightly between 1--2/1/3 of proximal part of leaf, with
rounded to acuminate or retuse apex, adherent to the stem in the base, but
channeled and deflexed or erect in distal 2/3 of leaf; cells at mid leaf
thin-walled, subisodiametric to oblong 35--50(-58) x 33--43 \um, trigones
large and bulging; along margin 20--38 \um, with slightly to obviously
thickened external wall; cuticle finely striolate everywhere but more
markedly in proximal part or sometimes loosely developed. Oil-bodies of the grape-cluster type,
consisting of distinctly glistening, homogenous globules, (1--)2--6(--8) per cell., spheric (5--)6--15 \um in diameter
to ovoid 6--14 x 7--19 \um. Sexual condition dioicous. Perianth terminal on main axis,
immersed to emergent for 1/3 of its length, conical pluriplicate and
turbinate to the mouth, ca. 490 x 490 \um, perianth in distal part 1-stratose,
cells rectangular to obliquely rectangular, 38--75 x 10--13 \um, walls
thickened, yellowish colored, perianth in proximal part 2-stratose, cells
linear to oblong rectangular, 125--200 x 20--25 \um, walls moderately
thickened. Perigynium rhizogenous ventrally, 2 times longer than perianth;
bracts just below perianth, in 2 pairs (due to high and robust perigynium),
ovate shortly 2-lobed, 1250--1400 x 1250--1300 (0.95--1.1:1). Androecia intercalary in 8--13 pairs
of bracts, with 1--2 antheridia per bract, stalk 2-seriate, ca. 63 x 18 \um;
bracts slightly inflated in base, but flattened and deflexed away the stem
above, purplish to deep-green purple in inflated area, more or less similar
in shape to sterile leaves, but complicated on a micro-slide due to inflation
in the base. Capsule with outer
cells 30--58 x 22--34 \um, each cell with 2--4 nodular thickenings on each
vertical wall and with 0--2 on horizontal wall; inner cells 40--110 x 10--22
\um. Elaters 8--12 \um in width,
2-spiral. Spores reddish to
golden-brown, 14--18 \um in diameter. On wet rocks in
streams, mostly in coniferous forest and crooked forests in oceanic regions;
0--1200 m; B.C.; Alaska, Wash. Endemic. 7. Plectocolea subelliptica (Lindberg ex Heeg) A. Evans in H.
Buch, A. Evans & Verdoorn, Ann. Bryol., 10: 42.
1938 Nardia subelliptica Lindberg ex Heeg, Verh. K.K. Zool.-Bot. Ges. Wien 43(Abh. 1): 69. 1893; Jungermannia subelliptica (Lindberg ex Heeg) Levier; Plectocolea obovata var. minor (Carrington) Schljakov; Solenostoma subellipticum (Lindberg ex Heeg) R.M. SchusterPlants 3--5 x 0.6--1
mm, erect to (rarely) ascending, brownish green, yellowish brown,
sepia-brown; fertile plants larger than the sterile. Stem 210--280 \um in width and 200--270 \um in height; dorsal
surface cells 43--75 x 10--13 \um, long-rectangular to linear, cuticle
distinctly striolate-papillose; in the stem cross section thick-walled, with
the exception of external wall of outer layer(!), walls brownish to colorless
in inner portion, trigones small, concave, outer cells ca. 30--40 \um in
diameter. Rhizoids reddish to
purplish and brown to nearly colorless. Leaves
inserted at 40--80° with axis, dorsally shortly decurrent ca. 1/5 of the stem
width, ventrally inserted subtransversely, not decurrent; 525--735 x 525--875
\um (1: 0.85--1), suborbicular to widely ovate and
ovate-lingulate, widest near the middle or at proximal 1/3, distinctly, but
not strongly concave-channeled; cells at mid leaf thin-walled,
subisodiametric, 22--30 x 22--28 \um, walls colorless, trigones distinct,
convex; along margin 20--25 \um, thin-walled, walls colorless to brownish,
trigones triangular to convex, cuticle striolate in proximal 1/3 of the leaf.
Oil-bodies (2--)3--5(--6)
per cell, 4--5(--5.5) x 6--10 \um, granulate. Sexual condition dioicous
(some innovations with 2 male generations divided by some pairs of sterile
leaves---heteroicous plants). Perianth terminal on main axis, conical, with ca. 8 archegonia
per gynoecium. Perigynium 1--1.3
of the perianth length; bracts just below perianth, in 1--2 pairs, ca. 1000 x
950 \um, wider in proximal 1/3, frequently with undulate margin. Androecia below perianth, commonly
divided from the latter by 1--3 pairs of sterile leaves, spicate, in 2--3
pairs of bracts, ca. 630--805 x 810--1050 \um, suborbicular
to orbicular triangle, wider in proximal 1/3 of the leaf, strongly inflated
near the base, purple near the inflated area, bract margin incurved, with 2
antheridia per bract, body grayish to almost colorless, stalk 2-seriate, ca.
87--100 x 15--20 \um. Capsule 500--1000
μm, ellipsoidal, capsule wall 2-stratose, outer
cell subquadrate to rectangular, 20--45 x 26--35 μm
with 2--3 nodular thickenings in each vertical wall; inner cells irregularly
linear, 30--90 x 8--20 μm with 6--18 annular
thickenings. Seta 100-- 150 μm in diameter and 1500--9000 μm
in length. Spores finely
papillose, spheric, 18--24 μm. Elaters 120--150 x 9--12, 2-spiral, with narrowed homogenous
ends. On moist soil and
rocks near stream, almost exclusively in mountains tundra or slightly
descending to forest belt along streams; 0--600; Greenland; B.C., N.S., Que.;
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