Tortella tortuosa var. tortuosa

Treatment appears in FNA Volume 27. Treatment on page 505. Mentioned on page 504, 506.

Plants green or yellow-green, appearing brown proximally, loosely foliose, some leaf-bases exposed, only slightly comose at stem apex. Stems conspicuously tomentose. Leaves in tight, complex spirals when dry, appearing soft or lax throughout the stem length, not fragile or erose, leaf tips nearly all present; conspicuously undulate, fragile or not, erect to erect-spreading when wet; undulate or plane; proximal laminal cells thin-walled and hyaline, sharply differentiated in shape and size from the papillose distal cells, which papillose in the area of contact; leaves keeled in section, costa in apical region exposed adaxially by up to two stereid cells in width or completely covered with an adaxial epidermis of quadrate papillose cells; leaf cross-section in one layer beside the costa, the lamina uniformly 1-stratose, lamina intact, the costa always differentiated into guide cells, stereids and epidermal-cells, adaxial stereid layer never disappearing toward the apex, adaxial epidermal layer typically absent apically in a medial groove to two stereid cells in width.


Phenology: Capsules mature late spring–summer (Jun–Aug).
Habitat: Calcareous regions, exposed or forest-shaded rock crevices, boulders, ledges of mountains or low, peaty soil and rotten wood, dry wooded hillsides or wet areas such as Thuja swamps, banks of streams over humus, river margins, northern regions in wet tundra and solifluction lobes
Elevation: low to high elevations (100-3800 m)

Distribution

V27 726-distribution-map.gif

Greenland, Alta., B.C., N.B., Nfld. and Labr. (Nfld.), N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon, Alaska, Calif., Colo., Idaho, Ill., Ind., Iowa, Maine, Mass., Mich., Minn., Mont., Nebr., Nev., N.H., N.Mex., N.Y., N.C., Ohio, Oreg., Pa., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., Wis., Wyo., Mexico, Central America (Guatemala), Europe, Asia, n Africa, Atlantic Islands (Iceland)

Discussion

Variety tortuosa is striking in its crisped leaves in spirals when dry, which, together with the strongly undulate margins, is a trait absent in other species in the genus or related genera. The long, multicellular, vitreous awn confluent with the lamina is also distinctive. Only in the case of very depauperate forms is var. tortuosa confused with Tortella fragilis, which has leaves more rigid, more or less erect, not or only slightly contorted when dry and regularly fragile with propaguloid modifications in the leaf apex. Occasional specimens of var. tortuosa have large laminal cells, on average 12 µm wide and attaining 14 µm in individual cells. The leaves never have incurved margins nor are they apically cucullate as in T. inclinata or in the younger leaves of T. rigens. Throughout the range of var. tortuosa occasional puzzling specimens may be found with cell sizes to 14 µm in the distal region of the leaves (Nebraska, Alberta, Arctic specimens). Absence of adaxial epidermal cells on the costa throughout much of the distal portion of the leaf may suggest T. inclinata in the loose sense, but epidermal cells may be found at least in the mid-proximal regions of most leaves. Occasional specimens determined as T. fragilis are richly perichaetiate specimens of T. tortuosa with extremely long perichaetial leaves throughout the stem resembling the subulate propagula at the leaf apices of the former species. The tips of these leaves, however, are not swollen at the distal ends.

Variety tortuosa exceptionally has a distinct central strand but this trait is generally absent. Trichostomum tenuirostre, with leaves also spirally twisted when dry, has hyaline proximal cells that extend indistinctly only a short way up the margin by a few cells in width, and the stem is not radiculose. Trichostomum tenuirostre usually has a large and distinct stem central strand, and the apex of the leaf often has low and distant teeth.

Selected References

None.

Lower Taxa

None.
elongate +  and quadrate +
not vesiculose +
of 1-2 rows +  and differentiated +
long-excurrent +  and short-excurrent +
Concave (?) +, Cucullate (?) +, Obtuse (?) +  and Acute (?) +
Patricia M. Eckel +
(Hedwig) Limpricht +
reniform +  and rounded +
papillose +  and smooth +
usually slightly wider +
somewhat broader +
Tortula tortuosa +
obovoid +
cucullate +
yellow +  and reddish-brown darker red or brown +
cleistocarpous +  and stegocarpous +
inclined +  and erect +
elliptic;cylindric +
0.15 cm1.5 mm <br />0.0015 m <br /> (0.33 cm3.3 mm <br />0.0033 m <br />) +
monoicous +, dioicous +  and sexual +
flat to bulging +
thick-walled +
green +  and clear +
0 (?) +  and 2 (?) +
papillose +
rhomboidal +, elongate +  and quadrate +
tomentose +  and bare +
excurrent +  and thin-walled +
quadrate +
differentiated +
7um +  and 10um +
autoicous +, dioicous +  and sexual +
setaceous-subulate +
Greenland +, Alta. +, B.C. +, N.B. +, Nfld. and Labr. (Nfld.) +, N.W.T. +, N.S. +, Nunavut +, Ont. +, Que. +, Sask. +, Yukon +, Alaska +, Calif. +, Colo. +, Idaho +, Ill. +, Ind. +, Iowa +, Maine +, Mass. +, Mich. +, Minn. +, Mont. +, Nebr. +, Nev. +, N.H. +, N.Mex. +, N.Y. +, N.C. +, Ohio +, Oreg. +, Pa. +, S.Dak. +, Tenn. +, Tex. +, Utah +, Vt. +, Va. +, Wash. +, Wis. +, Wyo. +, Mexico +, Central America (Guatemala) +, Europe +, Asia +, n Africa +  and Atlantic Islands (Iceland) +
low to high elevations (100-3800 m) +
differentiated +
multicellular +
conspicuous +
extensive +
Calcareous regions, exposed or forest-shadCalcareous regions, exposed or forest-shaded rock crevices, boulders, ledges of mountains or low, peaty soil and rotten wood, dry wooded hillsides or wet areas such as Thuja swamps, banks of streams over humus, river margins, northern regions in wet tundra and solifluction lobesgions in wet tundra and solifluction lobes +
yellow +  and orange-red or red +
disappearing +
0.15 cm1.5 mm <br />0.0015 m <br /> (0.35 cm3.5 mm <br />0.0035 m <br />) +
concave;keeled;channeled;ovoid;lanceolate or lingulate +
elongated +  and short +
distinct +
reduced +
not +  and fragile +
erect;erect-spreading +
plane +  and undulate +
differentiated +
crenulate-papillose +
subulate-acuminate +  and undulate +
undifferentiated +
dioicous +  and sexual +
multicellular +
denticulate +
long-rostrate;conic +
0.15 cm1.5 mm <br />0.0015 m <br /> (0.2 cm2 mm <br />0.002 m <br />) +
photosynthetic +
compound +, simple +, hollow +  and solid +
c--shaped +
0.5 cm5 mm <br />0.005 m <br /> (0.55 cm5.5 mm <br />0.0055 m <br />) +
lateral +  and terminal +
as stalked buds +  and short-foliose +
lateral +  and terminal +
branched-spiculose +  and twisted +
red +, orange +  and yellow +
consisting of 16 mostly 2 times cleft spiculose striate or papillose lanceolate teeth , , , , +  and rudimentary +
Capsules mature late spring–summer (Jun–Aug). +
perichaetiate +
Laubm. Deutschl. +
filiform +
thick-walled +  and thin-walled +
brown +  and hyaline +
long-rectangular +
straight +
2.7 cm27 mm <br />0.027 m <br /> (3.5 cm35 mm <br />0.035 m <br />) +
reddish;yellow +
elongate +
0.9 cm9 mm <br />0.009 m <br /> (2.7 cm27 mm <br />0.027 m <br />) +
2 +  and 1 +
not fragile +
yellowish-brown +
spreading;recurved +
contorted +  and crisped +
eradiculose +
rounded-triangular +  and rounded-pentagonal +
1 cm10 mm <br />0.01 m <br /> (6 cm60 mm <br />0.06 m <br />) +
Tortella tortuosa var. tortuosa +
Tortella tortuosa +
variety +
ovoid;cylindric +
0.65 cm6.5 mm <br />0.0065 m <br /> (0.7 cm7 mm <br />0.007 m <br />) +
flexuose +
widespreading +
linear-lanceolate +  and broadly narrowly concave or nearly plane +
0.3 cm3 mm <br />0.003 m <br /> (0.65 cm6.5 mm <br />0.0065 m <br />) +
orange +  and yellow +
perichaetiate +  and perigoniate +
yellow-green;green +
small +  and medium-sized +
0.11 cm1.1 mm <br />0.0011 m <br /> (0.14 cm1.4 mm <br />0.0014 m <br />) +