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BFNA Title: Imbribryum |
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XX. IMBRIBRYUM
Pedersen, Bryologist
108: 449. 2005 * [Greek imbri,
imbricate, and bryum, a moss, alluding to strongly imbricate leaves] John R. Spence Plants medium-sized to robust, in open to dense turfs or
gregarious on soil or rock, green, yellow-green, red-green or red. Stems
0.5--3(--4) cm, evenly foliate, innovations and fertile stem leaves somewhat
differentiated, leaves of innovations smaller, stems sometimes julaceous;
rhizoids sparse, at base of stem, occasionally in clusters along stem. Leaves
(0.6--)1.5--2.5(--3) mm, strongly imbricate
when dry, erect when wet, flat or concave, rarely decurrent; apex
obtuse to acuminate, apiculus absent; margins plane to strongly revolute,
smooth to serrulate distally, limbidium absent or occasionally present, of
1--2 rows of more elongate cells; costa percurrent to short-excurrent as an
awn, in cross-section generally with 1 layer of guide cells; distal and
mid-laminal cells vermicular to rhomboidal or hexagonal, 6--16(--22) \um
wide, mostly 3--6(--8):1, thin-walled to incrassate, rarely porose, proximal
laminal cells abruptly quadrate or rectangular, 1--2(--3):1, wider and
shorter than cells above, 1--2 layers of enlarged colored cells sometimes
present across leaf base, inflated group of pink sub-alar cells absent. Specialized
asexual reproduction by rhizoidal tubers, spherical, (100--)150--350 \um,
red, red-brown, orange, or pink, on rhizoids in proximal leaf axils. Sexual condition dioicous;
perigonial and perichaetial leaves not much differentiated, outer
perichaetial leaves larger, inner leaves smaller than leaves of innovations. Seta
long-exserted, stout to slender, straight or somewhat flexuose. Capsule
2--6 mm, pyriform to clavate, apophysis slender, not thickened, inflated or
rugose; inclined or nutant; peristome double, well developed, exostome teeth yellowish,
often separate at base, endostome high, not adherent to exostome, segments
well developed, with ovate perforations, cilia long and appendiculate or
sometimes nodose; operculum low-conic to convex, sometimes apiculate, rarely
rostrate. Spores small, smooth to papillose, 8--18(--20) \um. Species ca. 40 (6 in the flora): Nearly worldwide in
tropical, subtropical, temperate to boreal regions, rare in Arctic-alpine
climates, absent from Imbribryum consists of relatively large species
with elongate, evenly foliate stems and strongly imbricate leaves. Species
commonly grow on soil or rock, often associated with waterfalls, seeps and
springs, or splashed rock. The genus is especially well represented in the
mountains of SELECTED
REFERENCES Whitehouse, H. L. K. 1963. Bryum
riparium 1. Leaves
rigid, strongly imbricate when dry; distal laminal cells long, incrassate and
vermicular, more than 6:1. 2.
Costa percurrent to short-excurrent; limbidium absent; proximal laminal cells
gradually wider proximally, short-rectangular to quadrate; capsule pyriform
with short neck . . . 1. Imbribryum alpinum 2.
Costa
short to moderately long-excurrent into a slender awn; weak to moderately strong limbidium
present; proximal laminal cells abruptly enlarged, somewhat bulging,
rectangular; capsule with distinct, long neck. . . 3. Imbribryum microchaeton 1.
Leaves rigid to somewhat loosely imbricate; distal lamina cells shorter, not
vermicular, mostly 3--5:1. 3.
Stems strongly julaceous; leaves red or purple, rarely green, strongly
concave, apices rounded-obtuse to broadly acute, cucullate; costa percurrent; distal lamina cells
incrassate, oblique to costa, proximal laminal cells abruptly enlarged to inflated in 1--2
rows . . . 5. Imbribryum miniatum 3. Stems not or weakly julaceous; leaves red to
green or yellow, concave to flat, apices obtuse, acute, or acuminate, not
cucullate; costa not reaching apex to excurrent; distal lamina cells thin to
somewhat firm-walled, parallel to costa, cells at leaf base not abruptly
enlarged or inflated. 4. Leaves flat or weakly concave,
ovate-lanceolate, apices acute to acuminate; costa short-excurrent in
slender, sometimes denticulate awn . .
. 4. Imbribryum mildeanum 4. Leaves distinctly concave, ovate, apices
broadly acute or obtuse; costa not reaching apex or rarely percurrent, awn
absent. 5. Leaves green to yellow-green, lacking red tints,
loosely set, somewhat distant proximally along stem; proximal laminal cells
short-rectangular . . . 2. Imbribryum gemmiparum 5. Leaves red or red-green, red tints usually present, rigid
and imbricate, crowded; proximal laminal cells quadrate . . . 6. Imbribryum muehlenbeckii 1. Imbribryum alpinum ( Bryum alpinum Plants medium-sized, in open to dense turfs,
dark red to red-green, rarely entirely green, sometimes becoming black with
age. Stems 0.5--2(--3) cm, evenly foliate, somewhat julaceous, with
metallic sheen, older portions of stem sometimes densely radiculose. Leaves
1--3 mm, red to red green or sometimes green, strongly imbricate when dry,
erect when wet, rigid, broadly lanceolate to narrowly ovate or lingulate or
occasionally narrowly triangular, weakly concave, not decurrent; apex acute;
margins plane distally, revolute proximally, smooth to finely serrulate
distally, limbidium absent; costa reddish, percurrent to short-excurrent as a
stout awn; distal and mid-laminal cells elongate-hexagonal to vermicular,
(50--)60--80 x (8--)10--12 \um, mostly 6--8:1, strongly incrassate, parallel
to costa, proximal laminal cells abruptly quadrate to short-rectangular,
1--2(--3):1, occasionally a single row of colored cell present across leaf
base. Specialized asexual reproduction of red to red-brown spherical
rhizoidal tubers on rhizoids arising from leaf axils, 100--300 \um, scarce. Seta
1--3 cm, stout, more or less straight, red or red-brown to purple. Capsule
3--5 mm, pyriform, short-necked, inclined to nutant, red-purple; operculum
convex, apiculate. Spores smooth to papillose, yellowish, 10--15 \um. Capsules rare,
maturing spring--summer. Local and uncommon on moist acidic rock or soil over
rock; 0--2500 m; Greenland; Nfld. & Labr.; Alaska (St. Lawrence I.),
Calif., Colo., Ill., N.H.; w,n Eurasia; n Africa. Imbribryum alpinum is a relatively rare boreal-temperate species characterized by
narrow, shiny-metallic, red leaves and incrassate, elongate distal laminal
cells. Most reddish plants similar to
this species in the western 2. Imbribryum gemmiparum (De Notaris) J. R. Spence, Phytologia 89: 112.
2007 Bryum gemmiparum De Notaris, Comm. Soc. Crittog. Ital.2: 212
[112] [Cronaca Bryol. Ital. 1: 25] 1866 Plants medium-sized, in open to dense turfs, bright green to yellow-green
near tips, becoming stramineous with age. Stems 1--2(--3) cm, evenly
foliate, weakly julaceous, with or without metallic sheen, older portions of
stem sometimes densely brown-radiculose. Leaves 1--3 mm, green,
younger leaves sometimes yellowish, strongly imbricate when dry, erect when
wet, soft, strongly concave, weakly decurrent; apex rounded to acute; margins
plane throughout, rarely revolute
proximally, smooth, limbidium absent; costa not reaching apex or percurrent,
brown to yellow-brown; distal and mid-laminal
cells
hexagonal, (30--)40--60 x (14--)16--26 \um wide, mostly 3(--4):1,
thin-walled, proximal laminal cells abruptly short-rectangular, 2--3:1,
occasionally with quadrate cells intermixed, single row of colored cell
across leaf base absent. Specialized asexual reproduction of pink to
orange spherical rhizoidal tubers on rhizoids arising from leaf axils,
100--200 \um, rare [leafy bulbils in leaf axils]. Seta 1--3 cm, stout,
more or less straight, brown. Capsule 2--3 mm, pyriform, short-necked,
inclined to nutant, brown; operculum convex, weakly apiculate. Spores
small, smooth to papillose, yellowish, 12--18 \um. Capsules
rare, maturing spring--summer. Damp to wet calcareous soil or soil over rock,
often associated with springs; 0--1800 m; Alta., B.C., Ont.; Alaska, Ariz., Calif., Colo., Idaho, Maine, Mass., Mich., Minn., Mo.,
Mont., Mo., Nebr., Nev., N.J., N.Mex., N.Y., Okla., Oreg., Pa., S.Dak., Utah, Wash, Wyo.; s Europe; Asia
(Turkey); n Africa. European material of Imbribryum gemmiparum is reported to
produce small, leafy bulbils in leaf axils, but there is some controversy
over this as they are also sometimes described as short innovations with
stalks. Molecular studies show that at least some European material appears
to be related to the Gemmabryum dichotomum complex. However, North American material is much
larger in size, and is otherwise morphologically closest to other Imbribryum
species. Future studies may indicate that there are two species present, one
in North America and the second in Europe and the 3. Imbribryum microchaeton (Hampe) J. R. Spence, Phytologia 89: 112. 2007 Bryum microchaeton Hampe, Ann. Sci. Nat. Bot., sér. 5, 4: 342. 1865 Plants medium-sized, in open to dense turfs, pale shiny green to
brown-green. Stems 0.5--2(--3) cm, evenly foliate, not julaceous, with
metallic sheen, older portions of stem sometimes densely radiculose. Leaves 1--3 mm, pale shiny green,
becoming brown with age, strongly
imbricate when dry, erect when wet, rigid, weakly concave or flat, not
decurrent; apex acute to acuminate; margins strongly revolute throughout,
sometimes plane distally, smooth to finely serrulate, limbidium present, of
1--2 rows of elongate cells; costa short to moderately long-excurrent into a
slender awn; distal and mid-laminal
cells
vermicular to elongate hexagonal, (8--)10--12 \um wide, mostly 6--8:1,
strongly incrassate, proximal laminal cells abruptly enlarged, rectangular,
2--3:1, walls somewhat bulging, row of colored cells lacking across leaf
base. Specialized asexual reproduction by spherical rhizoidal tubers,
100--200 \um, red or red-brown, on rhizoids on stem. Seta 1--3 cm,
stout, more or less straight, brown to red-brown. Capsule 4--6, mm,
elongate-clavate, with long distinct neck, inclined to nutant, brown;
operculum convex, apiculate. Spores papillose, yellowish, 16--20 \um. Capsules
mature spring--summer. Rare on damp calcareous rock or soil over rock in
protected sites; 0--500 m; Imbribryum
microchaeton was recently collected in the 4. Imbribryum mildeanum (Juratzka) J. R. Spence, Phytologia 89: 112. 2007 Bryum mildeanum Juratzka, Verh. Zool.-Bot. Ges. Wien. 12: 967.
1862 Plants small to medium-sized, in open to dense turfs, bright green, older
portions of stems becoming stramineus. Stems 0.5--2(--3) cm, evenly
foliate, not julaceous, lacking metallic sheen, sometimes densely radiculose.
Leaves 1--3 mm, strongly imbricate when dry, erect when wet, rigid,
ovate-lanceolate, weakly concave or flat, not decurrent; apex acute to
acuminate; margins plane distally, revolute proximally, smooth to finely
serrulate distally, limbidium absent; costa short-excurrent in slender,
sometimes denticulate awn, brown, sometimes red proximally; distal and mid-laminal cells elongate-hexagonal,
(30--)40--60 x (8) 10--12 \um, mostly 4--6:1, thin to firm-walled, proximal
laminal cells abruptly quadrate, with a few short-rectangular cells
intermixed, 1--2:1, row of colored cells across leaf base absent. Specialized
asexual reproduction absent. Sporophytes unknown. Damp
soil along stream or in wetlands; 100--3500 m; Imbribryum
mildeanum is a small, inconspicuous
boreal-montane species that has been found in 5. Imbribryum miniatum (Lesquereux) J. R. Spence, Phytologia 89: 112.
2007 Bryum miniatum Lesquereux, Mem. Plants medium-sized, in open to dense turfs, dark red to red-green, rarely
green, sometimes purple-black. Stems 2--4(--5) cm, evenly foliate,
strongly julaceous, with metallic sheen, older portions of stem sometimes
densely radiculose. Leaves (1--)2--3 mm, purple-red to brown-red,
rarely dull olive-green, strongly imbricate when dry, erect when wet, rigid,
ovate, strongly concave, not or weakly decurrent; apex rounded-obtuse to broadly
acute, cucullate; margins plane throughout or rarely revolute proximally,
smooth to finely serrulate distally, limbidium absent; costa percurrent,
red-brown to brown; distal and mid-laminal
cells
elongate-rhomboidal, (40--)50--60 x (10--)12--14 \um, mostly 4--6:1, strongly
incrassate, arranged in rows oblique to the costa at 30--45° angle, proximal
laminal cells abruptly
enlarged to inflated, quadrate, with a few short-rectangular cells
intermixed, 1--2:1, with 2--3 rows of somewhat inflated colored cells present
across leaf base. Specialized asexual reproduction absent. Seta
1--3 cm, stout, more or less straight to flexuose, red or red-brown to
purple. Capsule 2--4 (5), mm, pyriform, inclined to nutant,
red-purple; operculum convex, apiculate. Spores smooth to papillose,
yellow-brown, 14--18 \um. Capsules
common, maturing spring--summer. Common on damp to wet siliceous rock or soil
over rock, often associated with waterfalls or springs; 0--1500 m; B.C.,
Ont.; Nfld. & Labr.; Ark., Calif., Idaho, Mo., Mont., Oreg., Pa., Wash.;
Atlantic Islands (Faroes). Imbribryum miniatum is a distinctive species, with
dark red-purple, julaceous stems, oblique and strongly incrassate distal
laminal cells, and colored inflated cells across the leaf base. Some
collections from 6. Imbribryum muehlenbeckii (Bruch & Schimper) Pedersen, Bryologist 108:
449. 2005 Bryum muehlenbeckii Bruch & Schimper, Bryol. Eur. 4: 163. pl.
381. 1846 Plants small to medium-sized, in open to dense turfs, dark red to red-green,
rarely entirely green. Stems 0.5--2(--3) cm, evenly foliate,
julaceous, lacking metallic sheen, older portions of stem sometimes densely
radiculose. Leaves 1--2(--3) mm, dark red to red-green or sometimes
dark green, strongly imbricate when dry, erect when wet, rigid, ovate,
distinctly concave, not or weakly decurrent; apex rounded to broadly acute;
margins plane distally, revolute proximally, smooth to finely serrulate
distally, limbidium absent; costa reddish, not reaching leaf apex or rarely
percurrent; distal and mid-laminal
cells
hexagonal, (40--)50--70 x (16--)18--24 \um, mostly (2--)3--4:1, not
incrassate, parallel to costa, proximal laminal cells abruptly quadrate, with
a few short-rectangular cells intermixed, 1--2:1, occasionally a single row
of colored cells reaching across leaf base. Specialized asexual
reproduction by orange-red to red-brown, spherical tubers on rhizoids
arising from leaf axils, 100--200 \um, scarce. Seta 1--3 cm, stout,
more or less straight, red or red-brown to purple. Capsule 2--3 mm,
short pyriform, nutant, dark red-brown; operculum convex, apiculate. Spores
small, papillose, yellow-brown, 14--18 \um. Capsules
rare, mature summer (June--Aug). Damp shaded siliceous rock or soil in rock
crevices, montane; 500--3000 m; Greenland; B.C., Ont., Nfld. & Labr.;
Calif., Idaho, Maine, Mich., Mont., Nev., N.H., N.Mex., N.Y., Oreg., Pa.,
Tenn., Wash.; w Europe; Asia (Caucasus); Atlantic Islands (Madiera). Imbribryum muehlenbeckii is similar to small forms of |
