BFNA Title: Bartramia
Author: D. Griffin, III 
Date: November 3, 2003
Edit Level: R Brum+
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
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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.