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BFNA Title: Bartramia |
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Bartramia -
Bartramiaceae 4. BARTRAMIA Hedwig,
Sp. Musc. 164. 1801, conserved name * [for the Pennsylvania colonist, John
Bartram] Plants small to robust, in loose to dense,
soft or rigid tufts, dull green to glaucous, sometimes yellowish or yellowish
brown above, radiculose proximally. Stems erect, often forked, 1--15
cm, roundish in cross section, cortex of somewhat firm walled cells forming
an indistinct to distinct hyaloderm. Leaves erect-appressed to
spreading or circinate, occasionally flexuose when dry, spreading when moist,
slenderly lanceolate-subulate from a more or less sheathing, non plicate
base; lamina 2- or 3-stratose at margins or throughout; margins revolute at
shoulders, serrate to serrulate distally, teeth single or paired; costa
usually strong and abaxially prominent, sometimes obscure in distal acumen
and low in profile, subpercurrent to excurrent; distal laminal cells small,
subquadrate to oblong linear, firm-walled, prorulose on both surfaces; basal
cells elongate to rectangular or linear, thin-walled or infrequently thick walled
toward costa, pale, smooth. Sexual condition dioicous, autoicous or
synoicous; perigonia gemmiform; perichaetial leaves little differentiated,
typically less prorulose and with a laxer areolation than in stem leaves. Seta
elongate, rarely short, erect or sometimes curved, smooth. Capsule
usually inclined and asymmetric, less frequently erect and symmetric,
subglobose, oblique mouthed, furrowed or rarely wrinkled when dry; operculum
convex with a low, blunt beak; annulus not distinct; peristome double, single
or lacking; teeth reddish brown, without intermediate thickenings on back,
smooth or papillose, sometimes cleft above the middle; endostome yellowish,
sometimes lacking, segments keeled, cilia rudimentary or none. Spores reniform
to subspherical, papillose, 15--40 µm. Species 90 (7
in the flora). Worldwide (except Antarctica), and especially diverse in
montane tropics. Bartramia is generally recognized by the
linear-lanceolate leaves and sheathing base with a distal lamina that is 2-
or 3-stratose at the margins or throughout. Bartramia stricta lacks
the sheathing leaf base but otherwise accords with the generic concept. SELECTED
REFERENCES Crum, H. A. and L. E. Anderson 1981. Bartramiaceae. In: Mosses of
Eastern North America, vols 1--2. New York. Flowers, S. 1935. Bartramia. In: A. J. Grout, Moss Fl.
N. Amer. Vol. II, part 3, pp. 157--163. Newfane, Vermont. Flowers, S. 1973.
Mosses: Utah and the West. Brigham Young University Press. Provo. Fransén, S.
1995. A taxonomic revision of Neotropical Bartramia
section Vaginella C. Muell.
Lindbergia 20: 147-179. Ireland, R. R. 1982. Moss Flora of the Maritime
Provinces. Publ. Bot. No. 13, National Museums of Canada, Ottawa. Lawton, E.
1971. Moss Flora of the Pacific Northwest. Hattori Botanical Laboratory.
Nichinan. 1. Leaves
large, linear, 4--7 mm, not strict, base not sharply distinct; plants large,
to 10 cm. 2.
Seta short, equaling or slightly longer than capsule . . 1. Bartramia halleriana 2.
Seta many times longer than capsule . . . . . . . . . . . . . 2. Bartramia
pomiformis
1. Leaves
smaller, subulate, usually abruptly narrowed and divergent from a hyaline,
sheathing base (except B. stricta), usually strict, 2.5--5 mm;
plants small, to 5 cm. 3.
Leaf base not sheathing, not abruptly distinct . . . . . . 3. Bartramia
stricta 3.
Leaf base sheathing, hyaline, abruptly distinct from distal lamina. 4.
Costa in distal limb obscure; distal laminal cells 25--45 µm, prorulae low .
. . . . . . 4. Bartramia ithyphylla 4.
Costa in distal limb prominent; distal laminal cells 10--30 µm, prorulae
high. 5.
Shoulders delicate, often eroded; leaf tips fragile, usually broken . . . . .
. . . . 5. Bartramia potosica 5.
Shoulders relatively firm, not eroded; leaf tips mostly intact. 6.
Basal cells of leaf differentiated with thick walled cells toward costa and
thin-walled cells toward margin; leaf margins revolute distal to shoulders .
. . . . . . . 6. Bartramia brevifolia 6.
Basal cells of leaf thin-walled throughout; leaf margins typically plane
distal to shoulders . . . 7. Bartramia
subulata 1. Bartramia
halleriana Hedwig, Sp. Musc., p. 164. 1801 Plants robust, green or yellowish brown, in
tight to lax tufts. Stems to 15 cm. Leaves crisped when dry,
erect-spreading when moist, sometimes secund, narrowly lanceolate,
long-subulate, base laxly sheathing; margins revolute from shoulders to high
in the acumen, entire proximally, denticulate in shoulders, spinose-dentate
above; costa excurrent, rough abaxially in the distal portion, spinose in
excurrent tip; basal laminal cells thin-walled from costa to margin; median
and distal cells 5--12 × 4(--10) mm. Sexual condition
autoicous or synoicous. Seta curved, 2(--8) mm, often 2 or 3 per
perichaetium. Capsule subglobose to pyriform, asymmetrical, furrowed,
1.5(--2.5) mm, persisting on older sections of stems, somewhat hidden by
leaves; operculum conic convex; peristome double; teeth lanceolate,
500--550 µm, strongly transversely
barred, reddish brown, granulose proximally, papillose above; endostome with
high basal membrane, segments striate papillose, slightly shorter than teeth,
cilia present, weakly developed. Spores 15--24 µm, warty papillose. Capsules
mature June--Sept. Crevices of shaded cliffs and on rock outcrops in humid
forests; 200--500 m; Alta, B.C.; Europe; Asia; s South America; Australia
(Tasmania); Pacific Islands (Hawaii, New Zealand). Vegetatively,
this species is quite similar to robust forms of Bartramia pomiformis;
the short, curved seta equaling or slightly longer than the capsule, however,
is distinctive. 2. Bartramia
pomiformis Hedwig, Sp. Musc. 164. 1801 Bartramia crispa
Brid., Musc. Rec. 2(3): 131.1 f. 4. 1803; B.
circinnulata J. K. A. Müller &
Kindberg; Bartramia glauco-viridis J. K. A. Müller &
Kindberg Plants green to glaucous, occasionally
yellowish, in dense to lax tufts. Stems 0.5--8 cm. Leaves
flexuose to crisped when dry from an erect, scarcely sheathing base, laxly
erect to erect-spreading when moist, narrowly lanceolate to
linear-lanceolate, apex subulate; margins revolute from base to distal
acumen, entire at base, coarsely toothed distally, teeth paired; costa
excurrent, rough abaxially in the distal portion, spinose in excurrent tip;
basal laminal cells thin-walled throughout; median and distal cells 4--15 × 4--8 µm. Sexual condition autoicous or synoicous. Seta erect, flexuose, 5--25 mm. Capsule globose to ovoid,
asymmetrical, furrowed, 1.5--2.5 mm; operculum conic convex; peristome
double; teeth lanceolate, 375--450 µm, reddish-brown, granulose-papillose;
endostome with high basal membrane, segments finely seriate-papillose,
somewhat shorter than teeth, cilia present, weakly developed. Spores
20--26 µm, warty-papillose. Capsules
mature Feb.--Nov. Soil or rock, occasionally at base of trees or on downed
trunks, usually in moist shady forests; 0--1100 m; sw Greenland; B.C.,
Labr., N.B., Nfld., N.S.; Nunavut, Ont., Que., Sask.; Ala.,
Alaska, Ark., Calif., Conn., Ga., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine,
Md., Mass., Mich., Minn., Miss., Mo., Mont., Nebr., N.C., N.H., N.J., N.Y.,
N. Dak., Ohio, Okla., Oreg., Pa., S.C., S. Dak., Tenn., Vt., Va., Wash.,
W.Va., Wis. (reported from D.C., Del., R.I.); South America (Tierra del
Fuego); Europe; n,c, and e Asia; n Africa; Atlantic Islands (Madeira). The
combination of narrowly lanceolate to linear-lanceolate leaves, often crisped
or flexuose when dry, together with the elongate seta that typically lofts
the capsule above the foliage are distinctive characters and serve to
distinguish Bartramia pomiformis from the similar species, B. halleriana.
The latter is restricted to western Canada in the flora area. 3. Bartramia
stricta Bridel, Musc. Rec. 2(3): 132. 7 f. 5. 1803 Plants glaucous to brownish green, usually in
tight tufts. Stems rigid, 1--3 cm. Leaves stiffly
erect-appressed when dry, erect-spreading when moist, 2.5--4 mm, narrowly
lanceolate from a truncate, non sheathing base; margins recurved from near
base to distal acumen, entire proximally, serrulate distally; costa
relatively slender, ca. 50 µm wide,
not filling acumen, abaxially prominent, rough abaxially in the distal
portion, excurrent; basal laminal cells thin-walled throughout; median and
distal cells 12--20 × 4--6
µm. Sexual condition synoicous. Seta straight, 10--15
mm. Capsule erect, symmetrical, subglobose to ovoid, striate;
operculum conic convex; peristome single; teeth lanceolate, 250--300 µm, reddish brown, striolate papillose
proximally, smooth above. Spores subspherical, 22--26 µm, warty-papillose. Capsules
mature Jan.--Aug. Soil or rocks; 10--2300 m; Calif., N.Mex., Tex.; B.C.; w, c
and s Europe; Asia (Syria, Turkey); n
and c Africa; Australia. This
widespread Old World species occurs infrequently in the southwestern portion
of the flora area with an isolated occurrence in western Canada. The strict,
appressed leaves lacking shoulders and with relatively uniform areolation
from base to acumen aid in recognition. When fertile, the erect, symmetrical
capsules with a single but well developed peristome are diagnostic. Confusion
of this species and Anacolia laevisphaera may occur, especially
with sterile material. In the latter the tips of the distal leaves are often
divergent when dry, spreading recurved when moist, with the inner basal cells
of the leaf more differentiated than those of the basal angles compared with
basal areolation in Bartramia stricta. 4. Bartramia
ithyphylla Bridel, Musc. Rec. 2(3): 132. 1 f. 6. 1803 Bartramia ithyphylla
ssp. rigidula (Lindberg) Kindberg; Bartramia ithyphylla var. breviseta
(Lindberg) Kindberg Plants soft green to glaucous, in lax to
tight tufts. Stems 1--3(--5) cm. Leaves stiffly erect when dry,
distal leaves occasionally divergent, linear subulate from an erect clasping
base (base glistening white when dry), with well developed, firm shoulders;
margins plane, entire proximally, serrulate to serrate distally, distal teeth
often paired; costa excurrent in a slender tip, obscure in distal limb, low
conic in cross section, appearing to fill the acumen; basal laminal cells
thin-walled, distal cells 25--45 ×
5--7 µm, prorulae relatively low. Sexual
condition synoicous; perichaetial leaves somewhat longer than stem
leaves, to 6 mm, more strongly clasping. Seta 8--30 mm. Capsule
inclined to cernuous, subglobose to ovoid, asymmetrical, 1 mm, furrowed;
operculum low conic, beak blunt; peristome double; teeth lanceolate,
300--400 µm, reddish brown, strongly
transversely barred, finely papillose proximally, smooth above; segments
smooth, irregularly lanceolate, 1/2-2/3 the length of the teeth and somewhat
adherent to them, cilia absent or weakly developed. Spores 25--40 µm, coarsely papillose. Capsules
mature July--Nov. Soil or rocks; 0--3800 m.; Greenland; Alta, B.C., N.B.,
Nfld., Nunavut, Que., Yukon; Alaska, Calif., Colo., Idaho, Maine, Mich.,
Mont., Nev., N.H., Oreg., Utah, Wash., Wyo.; South America (Argentina); n and
c Europe; Asia (Taiwan); n and c Africa. Bartramia ithyphylla
is essentially an Arctic-alpine species with disjunct populations in the high
mountains of Africa and in austral South America. In the flora area it
frequents tundra and montane forest habitats with occasional occurrence at
moderate to low elevations at northern latitudes. The glistening white leaf
base is distinctive. The obscure costa in the distal limb and the elongate
distal laminal cells bearing low prorulae distinguish B. ithyphylla from
other small species of the genus in the flora area. 5. Bartramia
potosica Montagne, Ann. Sci. Nat. Bot. sér. 2, 9: 56. 1838 Plants glaucous or yellowish green, usually
in dense tufts. Stems 0.5--4 cm. Leaves erect-appressed to
laxly erect when dry, erect-spreading to spreading recurved or somewhat
secund when moist; tips fragile, frequently broken, 3--8 mm, narrowly
lanceolate subulate from a sheathing base, shoulders well developed but
delicate, often eroded; margins plane or weakly revolute, entire proximally,
serrulate distally, teeth frequently paired; costa excurrent in slenderly
serrulate tip, abaxially prominent, rough abaxially in the distal portion;
basal laminal cells thin-walled, distal cells 10--25 × 5--6 µm. Sexual
condition apparently dioicous; not producing sporophytes in flora area. Cliff crevices
and over thin soil on canyon ledges, occasionally at base of boulders;
2000--3300 m.; Ariz., Colo., N.Dak., N.Mex.; Mexico; Central America; South
America. This species
is widespread in mountains of the Neotropics, from Mexico to Argentina. The
fragile leaf tips and delicate, often eroded, shoulders aid in
identification. While most fertile collections from south of the flora area
are dioicous, S. Fransén (1995) has encountered several synoicous plants from
Mexico and Guatemala which he recognizes as Bartramia potosica ssp.
synoica Fransén. Gametangia have
yet to be found on plants from the flora area. 6. Bartramia
brevifolia Brid., Bryol. Univ. 2: 737. 1827 Plants glaucous or brownish green, usually in
dense tufts. Stems 0.5--3 cm. Leaves erect to erect-spreading
and somewhat flexuose when dry, spreading when moist, 2--5 mm, narrowly
lanceolate-subulate from a sheathing base, not particularly fragile, tips
usually intact, shoulders well developed, firm; margins usually strongly
revolute, entire proximally, serrulate distally, teeth often paired; costa
excurrent in serrulate tip, rough abaxially in the distal portion, not
filling the acumen; basal laminal cells thick-walled toward costa,
thin-walled toward margins, distal cells 8--25 ×
4--8 µm. Sexual condition
synoicous or dioicous. Seta straight to slightly arcuate, 6--16 mm. Capsule
globose to ovoid, asymmetrical, furrowed, 1.5--2.5 mm; operculum low convex,
beak blunt; peristome single or double; teeth lanceolate, 200-350 µm, reddish brown, strongly transversely
barred, finely papillose proximally, vertically striate above; endostome
absent or, if present, rudimentary and represented by a basal membrane,
occasionally with segments less than 1/3 the length of the teeth; cilia
absent. Spores subspherical, dark brown, 25--35(--40) µm, coarsely papillose. Capsules
mature Nov. Rock crevices and canyon ledges; 1800--4000 m.; Ariz., N. Mex.;
Mexico; West Indies (Dominican Republic); Central America (Costa Rica, Guatemala); South America (Bolivia,
Colombia, Ecuador, Peru, Venezuela). Bartramia brevifolia is recognized by the differentiated leaf
base having thick-walled cells toward the costa and thin-walled cells toward
the margins. Material collected in Arizona by Bartram and identified as
either Bartramia microstoma Mitt. or B. glauca
Lor. is B. brevifolia. The type of B.
microstoma is not included with the
Mitten herbarium at NY, but several collections of this species from Latin
America have been annotated by Fransén as B.
brevifolia. S. Fransén (1995)
reduced B. glauca to synonymy under B.
potosica. 7. Bartramia
subulata Bruch and Schimper, Bryol. Eur. 4: 53. 315. 1846 (fasc. 31
Mon. Suppl. 1.1.1) Plants glaucous or dark green, in dense to
lax tufts. Stems 0.5--1 cm. Leaves
erect-appressed when dry, erect and slightly spreading when moist; narrowly
lanceolate from an erect, sheathing base, 1.5--2.5 mm, shoulders well
developed, form; margins plane to weakly revolute and entire proximally,
plane and serrulate distal to shoulders; costa percurrent in moderately stout
tip, not filling the acumen, rough abaxially in the distal portion; basal
cells thin-walled, distal cells 12--25 ×
4--6 µm. Sexual condition
autoicous or synoicous. Seta 1--2 cm. Capsule globose to ovoid,
erect, symmetrical, furrowed, 1-1.5 mm; operculum conic, beak low, blunt;
peristome lacking. Spores reniform, 24--35 µm, coarsely papillose. Capsules
mature Aug.--Nov. Bare soil in montane tundra; 3500--4000 m; Colo; Europe;
Asia (China; India; Japan). Limited in the
flora area to the high mountains of Colorado, this diminutive species with
the erect, symmetrical capsules might be mistaken for Bartramia stricta. The latter
species, however, has a well developed exostome, the leaf base is not
sheathing and shoulders are lacking or, at best, weakly developed. Reports of
Bartramia breviseta Lindb. (given as B.
ithyphylla var. breviseta (Lindberg) Kindberg by some
authors) from high elevations in Colorado likely represent
misidentifications. In Bartramia breviseta the capsules are overtopped
by the perichaetial leaves (the seta is 1--3 mm) and the costa fills the
acumen. As presently understood, authentic Bartramia breviseta is
an Arctic-alpine species of the Old World. Its precise relationship to Bartramia subulata requires further study.
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