Tortula subulata

   Plants in tufts or small turfs to 0.5 cm high, dark-green but with a glaucous cast due to the long awns, never glossy. Leaves spreading when moist but deeply keeled and crispate when dry, oblong to spatulate, to 2.5 mm long, 3–4: 1. Median laminal cells pluripapillose with papillae usually not C-shaped, to 15 µm broad, in straight rows, with lumen/wall ratio 3–4: 1, quadrate with rounded to angular lumens and small corner thickenings. Basal cells rectangular with straight lateral walls, to 20 µm broad, about 10: 1, smooth, thin-walled. Suprabasal median cells usually somewhat smaller, less papillose and thinner walled than adjacent laminal cells. Costa occupying about 1/8 of leaf base, gradually tapering and excurrent in an hyaline and smooth awn which may be more than 1/2 as long as the subtending lamina. Abaxial cells of costa narrowly rectangular, to 12 µm, 4–10: 1, smooth, not pitted on lateral walls. Adaxial cells of costa similar to the adjacent laminal cells. Costa cross-section about 2 times as broad as thick, with a strong dorsal stereid band, a single layer of guide cells, and with a well-developed ventral epidermis. Margin entire throughout, recurved to near the apex. Axillary hairs to 150 µm, to 10 cells long, with no basal brown cells. Rhizoids red-brown, to 20 µm wide at base, smooth and monopodially branched. Stem with central strand, rather thick-walled inner cortical cells and several rows of thicker walled, smaller outer cortical cells.

   Dioicous. Perichaetial leaves little differentiated from vegetative leaves. Seta to 25 mm, red-brown, straight, smooth. Capsule erect or almost so, with the urn cylindric but slightly arcuate, to 3 mm, about 6: 1. Operculum conic-rostrate, 1/4–1/2 of urn length. Annulus differentiated but not revoluble. Peristome straight to slightly dextrorse, pale to pinkish, to 0.7 mm long, with almost no basal membrane, densely papillose throughout. Exothecial cells long rectangular (more than 10: 1) and thick-walled with lumen/wall ratio 1–2: 1, not in regular rows, to 15 µm wide. Spores to 10 µm, nearly smooth.

Vouchers: Lake Co.: Cobb Mountain east of Geyser Rock, Toren & Dearing 7085 (CAS); Mariposa Co.: Summit Road about 2 miles north of Fish Camp, Sierra National Forest, Norris 85353; Modoc Co.: Highway 89 at Bear Creek, Norris 77940; Santa Cruz Co.: Eagle Rock, Kellman 886 (CAS); Shasta Co.: Lost Creek near Boundary Spring, Lassen Volcanic National Park, Showers 2220 (UC); Siskiyou Co.: about 1 mile east of Copper Butte, Norris 50243; Tehama Co.; South Fork Beegum Creek about 1.5 miles west of Rat Trap Gap, Norris 56978; Tulare Co.: Forest Trail 31E18 at Silver Creek, Sequoia National Forest, Shevock 9082.

Literature: Harthill et al. 1979; Holmberg 1969; Kellman 2003; Koch 1950a, 1951e; Koch and Ikenberry 1954; Lawton 1971; Mishler 1978; Showers 1982; Spjut 1971; Steere et al. 1954; Toren 1977. As Barbula subulata Lesquereux 1868; Watson 1880. As Tortula angustata Howe 1897.

Illustration References: Malcolm et al. 2009 p. 157; Ignatov and Ignatova 2003; Lawton 1971; Smith 1978; Zander 1993.

Elevation by latitude plot for Tortula subulata
   in California