1A Capsules when dry and empty strongly constricted below mouth, with ribs separated by narrow furrows, collapsed at the constricted area of the urn; exothecial bands formed by four to six rows of cells, reaching the mouth or nearly so; exostome teeth remaining tightly fused in pairs in empty capsules, bordered by a hyaline halo. Distribution west coast of North America...U. crispa s.s.
1B. Capsules when dry and empty not or slightly constricted below mouth, with ribs separated by more or less broad furrows in the upper half of urn; exothecial bands formed by two to four (five) rows of cells, visibly separated from the mouth by a ring of small thin-walled cells; pairs of exostome teeth partially splitting in empty capsules, teeth not bordered by a hyaline halo...2
2A. Endostome segments incurved when dry, uniseriate with thickened and prominent transverse walls; all the cells of the exothecial bands hyaline with pale yellow thickened lateral walls; leaves markedly crisped when dry, abruptly narrowing from a concave base. Distribution - western and eastern North America...U. intermedia
2B. Endostome segments variably bent when dry, uniseriate or irregularly biseriate with thin transverse walls; cells of the exothecial bands evenly pale yellow, at least in the two central rows; leaves slightly or moderately crisped when dry, gradually narrowing from a plane to slightly concave base. Distribution - widespread throughout eastern North America...U. crispula
Leaves when dry: curved to tortuous, rarely circinate VS tortuous to circinate, rarely falcate-tortuous.
Leaf base: ± elliptical, not obviously concave, gradually narrowing into the lamina VS ± obovate, distinctly concave, abruptly narrowing into the lamina.
Sporophyte total length: 1.8-4.0mm VS 3.0-6.5mm.
Capsule dry before spore release: short cylindrical to ovoid or ± urceolate VS ellipsoid to cylindrical-ellipsoid, rarely long ovoid.
Capsule furrow width when dry and empty: ± broad, somewhat sinuous or irregular VS broad.
Endostome appearance: slender and fragile VS robust and ± persistent.
Endostome segment position when dry: incurved or erect or patent VS incurved.
Muscol. recent. suppl. 4: 112. [1818] 1819.
Plants to 3.3 cm high, usually strongly crisped, growing in dense, medium to large cushions, olivaceous to dark green above, brownish to reddish below. Stems abundantly branched. Rhizoids common at stem base, frequently also scattered along stems, brown-reddish, smooth.
Leaves mostly tortuose or circinate, occasionally falcate when dry, erect-patent to squarrose when wet, (1.3)1.4-3.7(3.8) X 0.3-0.9(1.0) mm; leaf lamina lanceolate to narrowly lanceolate, unistratose, variably keeled; leaf base wide, obovate, rounded or oblate, distinctly concave, abruptly narrowing into lamina. Apex acuminate, sometimes channelled or with one margin irregularly incurved. Margins weak and variably recurved on one or both sides of lamina.
Costa ending at some distance from apex. Median and upper leaf cells rounded to elliptic, (6)9-16(22) X (6)9-12(17) µm, with moderately incrassate cell walls, with one or two low and simple, frequently inconspicuous papillae; basal leaf cells long rectangular to linear, (15)25-45(74) X (5)7-11(14) µm, with incrassate, sinuous, sometimes porose cell walls; differentiated basal marginal cells in (5)7-16(20) rows, forming a broad marginal band along leaf base.
Goniautoicous.
Perichaetial leaves differentiated, clearly longer and with wider and more elongated base than vegetative leaves, apex acute to acuminate, and costa frequently percurrent. Vaginula naked or variably hairy, with hairs partially two- to four-seriate, papillose, hyaline.
Sporophyte (3.0)3.2-6.8(7.4) mm long. Seta (0.3)0.5-4.4(5.0) mm long. Capsule long exserted. Urn cylindrical to ellipsoid when dry and full of spores; clearly urceolate, strongly constricted below mouth when dry and empty, then with eight long and prominent ribs separated by narrow furrows, collapsed at the constricted area of urn; (0.5)0.8-1.7(2.0) mm long, yellowish brown to reddish brown with concolorous ribs. Neck usually long, 0.5-3.0 times as long as urn, gradually tapering into seta. Exothecial bands 4 or 5 (6) cells wide, almost as long as urn, reaching the mouth or obscurely separated from it by a thin ring, 8-33(38) µm tall, formed by 1-3(4) layers of small, thin-walled cells; differentiated cells of the band evenly yellow to pale orange, not concolorous with other exothecial cells.
Stomata located in urn base and upper part of neck. Peristome double. Prostome frequent, usually incomplete and low, more rarely well developed. Exostome of eight pairs of teeth remaining tightly fused even in old capsules, yellowish to orange, recurved and lying on the exothecium when dry; outer surface outer peristome layer (OPL) densely ornamented by thin papillae, weakly striate in the distal portion; inner surface (exostomial Principal Peristome Layer [PPL]) smooth or finely papillose, frequently with distinct trabeculae; PPL visible externally as a hyaline halo as OPL edges partially retract. Endostome of eight robust and persistent segments, occasionally with up to eight additional intermediate, sometimes shorter processes; principal segments c. two-thirds the length of teeth, long triangular to subulate, uniseriate with transversal walls variably incrassate, incurved when dry, translucent and shiny; external surface (endostomial PPL) smooth; internal surface inner peristome layer (IPL) smooth to finely papillose below, variably papillose above, but never opaque because of the papillae density.
Operculum plane-convex to conic, rostrate, yellowish without differentiated basal ring. Calyptra hairy, hairs one- to four- (to five-) seriate, exceeding the beak. Spores (12)14-23(25) µm in diameter.
Distribution: West coast of North America.
Bryol. Univ. 1: 793. 1827.
= Ulota ulophylla ssp. crispula (Bruch) Giacom.
Plants to 4.2 cm high, variably crisped, growing in dense, small to large cushions, olivaceous above, brown-reddish to dark brown below. Stems abundantly branched. Rhizoids abundant at stem base and frequently ascending along stems, brownreddish, smooth.
Leaves curved to tortuose, more rarely circinate when dry, erect-patent to squarroserecurved when wet, (0.7)1.2-2.8(3.0) X (0.1)0.3-0.6(0.7) mm long; leaf lamina lanceolate to narrowly lanceolate, unistratose, variably keeled; leaf base elliptical, sometimes oblong to obovate, slightly concave, usually gradually narrowing into lamina. Apex acuminate to long acuminate, frequently channelled or with one margin irregularl incurved. Margins weak and variably recurved on one or both sides.
Costa ending at some distance from apex, sometimes close to it. Median and upper leaf cells rounded to elliptic, (5)8-18(31) X (5)8-12(18) µm, with moderately incrassate walls, smooth or with one or two low and simple, frequently inconspicuous papillae; basal leaf cells long rectangular to linear, (11)25-40(60) 9 (5)6-10(13) µm with incrassate, sinuous, sometimes porose cell walls; differentiated basal marginal cells in (1)3-8(10) rows, forming a relatively narrow marginal band along leaf base.
Goniautoicous.
Perichaetial leaves somewhat differentiated, lanceolate to widely lanceolate, longer and wider than regular leaves, leaf base oblong to obovate, variably sheathing. Vaginula hairy, sometimes densely, with hairs partially two- or three-seriate, papillose, hyaline.
Sporophyte (1.8)2.0-3.8(4.0) mm long. Seta 0.9-2.1(3.0) mm long. Capsule exserted. Urn short cylindrical to ovoid or slightly urceolate when dry and full of spores; short cylindrical, somewhat urceolate, or obconic (gradually tapering from mouth to seta), not or slightly constricted below mouth when dry and empty, then with eight prominent ribs usually separated by broad, somewhat sinuous or irregular furrows; (0.6)0.7-1.2 mm long, pale yellow to brownish yellow, with darker ribs. Neck long, 0.4-2.0 times as long as urn, gradually tapering into seta. Exothecial bands two to four cells wide, almost as long as urn, separated from the mouth by a conspicuous ring, (12)18-65(75) µm tall, formed by (1)2-4(5) layers of small, thinwalled cells; differentiated cells of the band evenly pale yellow and not concolorous with other exothecial cells, rarely hyaline and concolorous (eastern North America).
Stomata located in urn base and upper part of neck. Peristome double. Prostome frequent, usually incomplete and low, sometimes developed up to one-third to one-half of teeth length. Exostome of eight pairs of teeth with marked tendency to split into 16, yellowish to pale orange, recurved and lying on the exothecium when dry; outer surface (OPL) ornamented by thick papillae, sometimes becoming longitudinally striate or with crests in upper one-half to one-third; inner surface (exostomial PPL) smooth, occasionally with sparse, sometimes aligned, papillae in upper half; PPL not externally visible at tooth margins. Endostome of eight slender and fragile segments, occasionally with up to eight additional intermediate, sometimes shorter processes; principal segments almost as long as teeth, almost linear with variably widened base, irregularly biseriate, with thin transversal walls, incurved, erect or patent when dry, variably translucent at base or lower half, more or less opaque in upper part; external surface (endostomial PPL) smooth; internal surface (IPL) with a dense reticulum of striae which, in extreme cases, may look like an opaque and almost smooth ornamentation.
Operculum plane to convex, rarely conic, rostrate, yellowish, usually with an orange to reddish basal ring. Calyptra hairy, hairs two- or three- (four-) seriate, variably exceeding the beak. Spores (18)20-29(30) µm in diameter.
Distribution: Amphi-Atlantic, widespread throughout eastern North America.
Syn. musc. Eur. (ed. 2): 305. 1876.
= Ulota ulophylla var. intermedia (Schimp.) Jansen & Wacht.
Plants to 3.5 cm high, crisped, growing in dense, medium to large cushions, olivaceous to dark green above, dark brown below. Stems abundantly branched. Rhizoids at stem and branch bases, sometimes ascending along stems, brown-reddish, smooth.
Leaves tortuose or circinate, occasionally falcate-tortuose when dry, erect-patent to squarroserecurved when wet, (1.0)1.5-3.4(3.7) X 0.3-0.7(0.8) mm; leaf lamina lanceolate to narrowly lanceolate, unistratose, variably keeled; leaf base obovate to orbicular, rarely oblate or elliptic, distinctly concave, abruptly narrowing into lamina. Apex acuminate to long acuminate, sometimes acute, frequently channelled. Margins weak and irregularly recurved on one or both sides, crenulate to crenulate-papillose.
Costa ending at some distance from apex. Median and upper leaf cells rounded to elliptic, 7-16(17) X (6)7-13 µm, with moderately incrassate walls, with one or two simple or bifurcate papillae; basal leaf cells long rectangular to linear, (18)20-63(68) X 5-10(12) µm, with incrassate, sinuous, sometimes porose cell walls; differentiated basal marginal cells in (2)3-14(16) rows, forming a broad marginal band along leaf base.
Goniautoicous.
Perichaetial leaves differentiated, clearly longer than regular leaves, leaf base long ellipsoidal to oblong-ellipsoidal, sometimes obovate or almost orbicular, frequently sheathing, apex acute to shortly acuminate. Vaginula naked or hairy, with hairs partially one- or two- (three-) seriate, papillose, hyaline.
Sporophyte (3.0)3.2-6.5 mm long. Seta 1.0-4.0(5.5) mm long. Capsule long exserted. Urn ellipsoidal to cylindrical-ellipsoidal, rarely long ovoid when dry and full of spores; cylindrical, oblongcylindrical, obconic or elongate-ovoid, not constricted below the mouth when dry and empty, then with prominent ribs separated by broad furrows; (0.6)0.8-1.5(1.7) mm long, pale yellow to brownish yellow, with darker ribs. Neck long, 0.5-2.0 times as long as urn, gradually tapering into seta. Exothecial bands two to four (five) cells wide, almost as long as urn, separated from the mouth by a conspicuous ring, (15)18-113(238) µm tall, formed by (one-) two to six (-seven) layers of small, thin-walled cells; differentiated cells of the band hyaline with pale yellow incrassate lateral walls, concolorous with other exothecial cells.
Stomata located in urn base and upper part of neck. Peristome double. Prostome frequent, incomplete and low or with fragments irregularly sparse along teeth, sometimes well developed. Exostome of eight teeth pairs with tendency to split into 16, pale yellow with bases sometimes becoming darker, brown-orange, recurved and lying on the exothecium, except sometimes at their bases, when dry; outer surface (OPL) with a dense papillose reticulum, sometimes the papillae fused into transverse striae in lower part or into longitudinal weak striae in upper third; inner surface (exostomial PPL) smooth or finely papillose, sometimes the papillae forming thin lines in lower half, frequently with marked trabeculae and fenestrate in upper third; PPL not visible externally at tooth margins. Endostome of eight robust and usually persistent segments, occasionally with up to eight additional intermediate, sometimes shorter processes; principal segments three-quarters as long as teeth, broadly linear with variably widened base to subulate, uniseriate, usually with incrassate and prominent transverse walls, incurved when dry, hyaline and usually shiny; external surface (endostomial PPL) smooth; internal surface (IPL) smooth or finely papillose in lower half, variably reticulate above but never opaque because of the ornamentation density.
Operculum plane to convex, sometimes conic, rostrate, yellowish without differentiated basal ring. Calyptra variably hairy, hairs (one-) two- to four- (five-) seriate, not or scarcely exceeding the beak. Spores (15)18-35(38) µm in diameter.
Distribution: Western and eastern North America.
from - "Integrative taxonomy sheds light on an old problem: the Ulota crispa complex" - Caparros et al 2016
Botanical Journal of the Linnean Society, Volume 180, Issue 4, April 2016, Pages 427-451, https://doi.org/10.1111/boj.12397
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