BFNA Title: Ulota
Ulota - Orthotrichaceae
8. Ulota C. Mohr, Ann. Bot. 2: 540 1806 * [referring to curled leaves of some species]
Dale H. Vitt
Plants to 5.5 mm. Stems erect or rarely creeping, ± branched. Leaves contorted-crisped to slightly twisted and erect-curved when dry, erect-spreading to spreading-flexuose when moist, 1--1.4 mm, oblong- to linear-lanceolate, usually from an ovate base, acuminate, acute, or narrowly obtuse, base usually clasping stem; margins entire, plane or reflexed; costa ending near apex or rarely excurrent; distal cells 6--13 µm wide, hexagonal-rounded to elliptic, with very incrassate walls and conical papillae, sometimes smooth; interior basal cells radiating from insertion, rectangular-elongate to elliptic, often nodose, very thick-walled, usually orange, sharply grading to thin-walled, hyaline, quadrate marginal cells. Brood bodies rarely present, if so, then only at tips of leaves. Perichaetial leaves larger. Sexual condition autoicous or rarely dioicous. Seta 1.5--10 mm. Capsule 0.6--3.2 mm, fusiform-cylindric, ovate-oblong or rarely obovate, lightly 8-plicate at mouth to moderately 8-ribbed entire length, fully exserted, mouth puckered or capsule ± constricted beneath mouth, gradually contracted to seta through long neck; stomates superficial; peristome double or single; exostome teeth 8, sometimes split to 16, ± perforate at tips, finely and densely papillose to papillose-striate, reflexed or erect and flexuose; endostome segments 8 or lacking, smooth, reticulate or papillose-striate, incurved; preperistome never present. Calyptra smooth, hairy, mitrate, short-conic to conic, not plicate, usually deeply split several times at base. Spores isomorphic.
Species ca. 60 (9 in the flora): temperate in
distribution; North America, South
America, Europe, Asia,
The genus Ulota
consists of approximately 60 species of which over half are restricted to the
Southern Hemisphere. Generally,
1. Plants dioicous, rarely fruiting; leaves with apical brood-bodies; maritime habitats . . 9. Ulota phyllantha
1. Plants autoicous, almost always fruiting; leaves without apical brood-bodies; non-maritime habitats.
2. Capsules distinctly smaller at mouth than at middle; peristome single; exostome teeth erect, flexuose.
3. Capsules fusiform, strongly 8-ribbed . . 5. Ulota drummondii
3. Capsules clavate or obovate, puckered and lightly 8-plicate at mouth . . 2. Ulota coarctata
2. Capsules constricted beneath mouth or evenly tapering to seta from mouth; peristome double; exostome teeth reflexed.
4. Perichaetial leaves clasping seta, rounded, smooth, ligulate . . 1. Ulota barclayi
4. Perichaetial leaves not differentiated from stem leaves.
5. Plants differentiated into creeping and upright stems; leaves with filiform-acuminate apex, linear-lanceolate; stomates near middle of capsule; spores 35--60 µm. . 7. Ulota megalospora
5. Plants with all branches upright; leaves with acute or obtuse apex, lanceolate or linear-lanceolate; stomates in neck of capsule; spores 16--29 µm.
6. Leaves straight, erect-appressed when dry, not crisped or contorted . . 6. Ulota hutchinsiae
6. Leaves flexuose, crisped, or contorted when dry.
7. Plants growing on rock; leaves flexuose but not contorted or twisted when dry . . 4. Ulota curvifolia
7. Plants growing on trees; leaves contorted, crisped, or twisted when dry.
8. Interior basal cells thin-walled, rectangular, not colored; calyptra sparsely hairy; leaves flexuose-crisped when dry
. . . Ulota barclayi
8. Interior basal cells thick-walled, elliptic-elongate, yellowish; calyptra densely hairy; leaves crisped-contorted when dry.
9. Plants less than or equal to 4 cm; seta usually equal to or greater than 4 mm; capsules widest at mouth and gradually tapering to seta; leaves contorted and tightly twisted around themselves; western North America . . 8. Ulota obtusiuscula
9. Plants less than or equal to 2.5 cm; seta always less than 3 mm; capsules constricted beneath mouth; leaves loosely twisted-crisped, not twisted around themselves; eastern North America . . 3. Ulota crispa
1. Ulota barclayi Mitten, J. Linn. Soc. London, Bot. 8: 26. 1864
Plants to 2.5 cm (usually less than 1 cm). Stem leaves slightly crisped and curved when dry, narrowly lanceolate to oblong-lanceolate, 1.5--2.5 mm, leaf base oblong; margin broadly reflexed or plane, distal laminal cells 8--12 µm wide, smooth or with small papillae; basal laminal cells hexagonal-rounded, thick-walled. Sexual condition autoicous. Seta 1.8--3.2 mm. Capsule oblong-ovate when mature, oblong-cylindric when old and dry, 1--2 mm, strongly 8-ribbed ± 3/4 length; stomates numerous in neck and proximal portion; peristome double; endostome segments 8, incurved, delicate, caducous; exostome teeth 8, ± split to 16, reflexed, evenly papillose. Calyptra short-conic, sparsely hairy or with few hairs near apex. Spores 16--21 µm.
Tree trunks and branches; at low elevations; B.C.;
barclayi, described from
2. Ulota coarctata (Palisot de Beauvois) Hammar, Mon. Orthotr. Ulota Suec. 25. 1852
Orthotrichum coarctatum Palisot de Beauvois, Prodr. 80. 1805; O. ludwigii Bridel; Ulota ludwigii (Bridel) Bridel
Plants 0.5--1.5 cm. Stem leaves slightly twisted, erect-curved when dry, narrowly lanceolate to lanceolate, 1.5--2.5 mm; leaf base ovate or oblong; margin reflexed; distal laminal cells 8--11 µm wide, with small conic papillae, sometimes almost smooth; basal laminal cells elongate-linear, thick-walled, sometimes ± nodose. Sexual condition autoicous. Seta 1.5--4 mm. Capsule obovate to oblong-obovoid, 1.1--2.8 mm, puckered, lightly 8-plicate at mouth, otherwise smooth; stomates numerous in neck; peristome single; no endostome segments; exostome teeth 8, splitting to 16, erect, densely and obscurely papillose. Calyptra oblong-conic, very hairy. Spores 19--26 µm.
Trunks and occasionally branches of deciduous trees, rarely on conifers; low elevations; N.B., Nfld., N.S., Ont., P.E.I., Que.; Conn., Idaho, Maine, Md., Mass., Mich., N.H., N.J., N.Y., N.C., Pa., Tenn., Vt., Va., W.Va., Wis.; Europe; Asia.
This rather uncommon species prefers climax beech-maple
forests, and is distributed from
3. Ulota crispa (Hedwig) Bridel, Mant. Musc. 112. 1819
Orthotrichum crispum Hedwig, Spec. Musc. 162: 1801; O. connectens Kindberg; Ulota crispula Bridel; U. connectens (Kindberg) Macoun & Kindberg
Plants to 2.5 cm. Stem leaves twisted-crisped when dry, narrowly-lanceolate to linear lanceolate, 1.7--3 mm; leaf base ovate, concave; margin plane to reflexed; distal laminal cells 7--10 µm wide, with conic papillae; basal laminal cells linear-elongate grading to quadrate at margin, thick-walled. Sexual condition autoicous. Seta 1--3 mm. Capsule short-oblong to oblong, 0.8--2 mm, strongly 8-ribbed from 3/4 to entire length; stomates numerous in neck and lower 1/3; peristome double; endostome segments 8, smooth; exostome teeth 8, reflexed, densely papillose below, lightly papillose-reticulate above. Calyptra conic, very hairy. Spores 23--29 µm.
Common on branches and trunks of trees, especially Betula species; low elevations; N.B., Nfld., N.S., Ont., PE.I., Que.; Conn., Del., Ga., Ill., Ky., Maine., Md., Mass., Mich., Minn., N.H., N.J., N.Y., N.C., Ohio., Pa., R.I., S.C., Tenn., Ver., Va., W.Va., Wis.; Europe; Asia.
This species is distributed from
4. Ulota curvifolia (Wahlenberg) Liljeblad, Utkast. Svensk. Fl. Ed. 3: 546. 1816
curvifolium Wahlenberg, Fl. Lapp. 365. 1812; O.
crispum var. curvifolium
(Wahlenberg) Wahlenberg; Ulota
Plants to 0.3 cm. Stem leaves flexuose-crisped and loosely appressed when dry, lanceolate, 1--2.2 mm, leaf base ovate, narrowly obtuse; margin reflexed, distal laminal cells 8--12 µm wide, with 1--2 large conic or forked papillae; basal laminal cells linear, grading to quadrate at margin, very thick-walled. Sexual condition autoicous. Seta 1.5--3 mm. Capsule ovate-oblong when mature, oblong to oblong-cylindric when old, 1.2--1.7 mm, strongly 8-ribbed for 3/4 length; stomates numerous in neck; peristome double; endostome segments 8, smooth; exostome teeth 8, incurved, smooth. Calyptra conic, very hairy. Spores 16--20 µm.
Acidic rock in montane and subarctic areas; 0--2000 m; Greenland; Alta., B.C., Lab., Man., Nfld., Nun., N.W.T., N.S. Ont., Que.; Alaska; n Europe; Asia.
This is a Canadian shield species in eastern North America, which has a range from the northern shore of Lake Huron and Lake Superior north to Hudson Bay and Baffin Island westward across the sub-Arctic of the Northwest Territories and southward in the Canadian Rockies to Alberta and southern British Columbia. This rock-growing species has strongly papillose, thick-walled distal leaf cells and stiffly flexuous leaves. Ulota hutchinsiae is more southern and has less papillose distal leaf cells and straight leaves. As with most other species of the genus, U. curvifolia has a rich chestnut brown coloration that helps indicate its identity in the field.
5. Ulota drummondii (Greville) Bridel, Bryol. Univ. 1: 299. 1826
Orthotrichum drummondii Greville, Scottt. Crypt. Fl.; Ulota bicolor Bridel; Ulota funstonii Grout
Plants 0.5--1.2 cm. Stem leaves flexuose to ±contorted-flexuose, loosely erect when dry, lanceolate, 1.7--3.7 mm, leaf base ovate; margin plane or slightly reflexed, distal laminal cells 7--11 µm wide, smooth or slightly papillose; basal laminal cells rounded to elliptic, thick-walled. Sexual condition autoicous. Seta 5--5.5 mm. Capsule cylindric and contracted at mouth when mature, fusiform to fusiform-cylindric when old and dry, 1.7--3.2 mm, 8-ribbed 1/2--2/3 length; stomates numerous in neck and basal portion; peristome single; no endostome segments; exostome teeth 8, splitting to 16, erect and flexuose-incurved, obscurely and densely papillose. Calyptra conic, very hairy. Spores 21--24 µm.
Twigs and trunks of conifer and
decidious trees, especially in dense coastal forests; low elevations; B.C.,
The distribution of this species is bicentric---it is known from both coasts in North America, from Newfoundland south to Cape Breton and the Gaspé in the East, and from along the Aleutians south to the Queen Charlotte Islands in the West. This species is distinguished by narrowly fusiform capsules that are ribbed almost their entire length and are somewhat smaller at the mouth. The whitish single peristome is erect and flexuose and easily distinguishes this species from U. crispa with a reflexed exostome. Additionally, the leaves are blunt and not strongly twisted-contorted.
6. Ulota hutchinsiae (Smith) Hammar, Mon. Orthogr. Ulot. Suec. 27. 1852
Orthotrichum hutchinsiae Smith, Engl. Bot. 36: 2523. 1813; Orthotrichum americanum Palisot de Beauvois
Plants 0.4--4 cm. Stem leaves erect-appresssed, stiff, ± slightly curled or twisted around stem, never contorted or crisped when dry, narrowly lanceolate-oblong to lanceolate, broadly acute, to obtuse, 1.4--2.2 mm, leaf base not much differentiated to broad, oblong; margin ±recurved to near apex, distal laminal cells 7--14 µm wide, with small to large, conic to clavate papillae; basal laminal cells rectangular to elongate-rectangular, ± nodose, thick-walled. Sexual condition autoicous. Seta 2.5--3.5 mm. Capsule oblong-elliptic to cylindric-fusiform, 1.1--2 mm, slightly to moderately 8-ribbed 1/3--1/2 length; stomates numerous in neck; peristome double; endostome segments 8, papillose-striate; exostome teeth 8, rarely splitting to 16, reflexed, densely papillose, perforate near tips. Calyptra oblong-conic, very hairy. Spores 9--16 µm.
Varieties 2 (2 in the flora): e and n North America; Europe;
The species is continuously distributed in eastern North
1. Leaves 1-stratose, distal leaf cells 8--14 µm wide, with non-incrassate walls and small, conic papillae. . . . 5a. Ulota hutchinsiae var. hutchinsiae
1. Leaves 2-stratose distally, distal leaf cells 7--10 µm wide, with very incrassate walls and large, clavate papillae. . . . 5b. Ulota hutchinsiae var. rufescens
6a. Ulota hutchinsiae (Smith) Hammar var. hutchinsiae
Stem leaves acute to narrowly obtuse; leaf base 1-stratose in distal portion; distal laminal cells 8--14 µm wide, with small, conic papillae; basal laminal cells ± nodose, thick-walled.
Acidic rocks; low to mid-elevations; N.B., Nfld., N.S., Ont., Que.; Ala., Alaska, Ariz., Ark., Conn., Del., Ga., Ky., Maine, Md., Mass., Mich., Minn., Mo., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., S.D., Tenn., Vt., Va., W.Va.; Europe; Asia.
Ulota hutchinsiae var. rufescens (E.