418 DICRANACEAE · Dicranum denticulate at apex; laminae 1-stratose or with some 2- acute; margins entire or serrulate at apex; laminae 1- stratose regions in patches between margin and costa; stratose or with 2-stratose regions near apex; costa long- costa excurrent, 1/5–1/4 the width of the leaves at base, excurrent, 1/6–1/4 the width of the leaves at base, abaxial smooth or slightly rough on abaxial surface in distal half, ridges absent; leaf cells smooth, with a row of guide cells, abaxial ridges absent, with a row of guide cells, two no stereid bands, 1 or 2 rows of cells above and below weakly developed stereid bands above and below, not guide cells with slightly thickened walls and large lumens, extending above the leaf middle, adaxial and abaxial the adaxial and abaxial epidermal layers of cells not epidermal layers of cells not differentiated or with a few differentiated; cell walls between lamina cells not bulging; cells enlarged in both layers; cell walls between lamina alar cells 1-stratose, differentiated, not extending to costa; cells not or weakly bulging; leaf cells usually smooth; proximal laminal cells rectangular to linear, with a few alar cells 1-stratose or with a few 2-stratose regions, well- pits, (46–)60–105(–120) × (4–)6–16(–22) µm; distal differentiated, often extending to costa; proximal laminal laminal cells quadrate, rounded or short-rectangular, not cells rectangular to short-rectangular, not pitted or with pitted, (12–)17–25(–32) × (8–)10–11(–12) µm. Sexual few pits, (11–)33–42(–51) × (7–)9–10(–13) µm; median condition dioicous; male plants as large as females; laminal cells regularly quadrate, not pitted, (9–)15–22 interior perichaetial leaves gradually narrowed to a (–26) × (5–)7–8(–11) µm; distal laminal cells small, subulate apex, convolute-sheathing. Seta 1.5–2.5 cm, quadrate, not pitted. Sexual condition dioicous; male solitary, yellow to light brown. Capsule 1.4–2.5 mm, plants as large as females; interior perichaetial leaves straight and erect, smooth, often irregularly wrinkled abruptly long-acuminate, convolute-sheathing. Seta 1– when dry, yellow to light brown; operculum 1–1.8 mm. 1.6 cm, solitary, yellow to reddish brown. Capsule 1.5– Spores 10–18 µm. 2.5 mm, straight, erect, smooth, slightly furrowed when Capsules mature in summer. Frequently on rotten logs, dry, brown to reddish brown; operculum 1–1.5 mm. stumps, or tree bases in woodlands, sometimes on humus Spores 9–22 µm. or humus over rock; 150–2200 m; Alta., B.C., N.W.T., Capsules mature in spring. Commonly growing on Sask.; Alaska, Calif., Colo., Idaho, Mont., Nebr., Oreg., the base of trees (usually deciduous but sometimes S.Dak., Utah, Wash., Wyo.; Europe. coniferous, especially Thuja), rotten logs, stumps, rarely Dicranum tauricum has been reported from Utah by soil and acidic or limestone rock; 40–1400 m; Man., N.B., S. Flowers (1973). It is a western North American species, Nfld. and Labr. (Nfld.), N.S., Ont., P.E.I., Que.; Alaska, occurring on trees and rotten wood. It is recognized as Ill., Ind., Iowa, Ky., Maine, Md., Mass., Mich., Minn., glossy plants with broken off leaf tips and straight, erect N.H., N.Y., N.C., Ohio, Pa., Tenn., Vt., Va., W.Va., Wis.; capsules. It can be confused with D. fragilifolium, which Europe; Asia. also has broken off leaf tips and whose range overlaps Dicranum viride has been reported from Alaska by with D. tauricum. For distinctions see discussion under I. A. Worley and Z. Iwatsuki (1970) and from Kentucky 21. D. fragilifolium. by J. A. Snider et al. (1988). It is a not uncommon species of the eastern United States and southeastern Canada. It 25. Dicranum montanum Hedwig, Sp. Musc. Frond., rarely produces sporophytes and is distinctive because 143. 1801 of the fragile, deciduous leaf tips, which probably serve Plants in dense tufts, usually with as a means of asexual reproduction. It has on occasion weak, clustered branchlets near been confused with 22. D. fulvum, which has nearly the 5 stem apices with small, linear, 5 same distribution, and rarely with 21. D. fragilifolium 5 erect-spreading, costate leaves,

5 where their ranges overlap in the Great Lakes region. 5 5 5 5 strongly crisped when dry, 5 5 5 5 5 For distinctions see the discussions under those species. 5 55 5 5 5 5 5 yellowish green to dark green, 5 5 5 55 5 5 5 5 5 5 5 5 5 5 5 5 5 dull. Stems 0.5–3(–5) cm, densely 24. Dicranum tauricum Sapjegin, Bot. Jahrb. Syst. 46: tomentose with white to reddish 10. 1911 brown rhizoids. Leaves erect- Dicranum strictum D. Mohr spreading, cirrate to strongly crisped when dry, smooth, Plants in dense tufts, light green (1–)2–3(–4) × 0.2–0.5 mm, concave below, subtubulose 5 to yellowish green, glossy. Stems to keeled above, lanceolate at base, acuminate above to 5 0.5–3 cm, tomentose below with an acute apex, rough above on abaxial surface; margins 5 5 5 irregularly serrate to serrulate in distal half of leaves; 5 whitish or reddish brown rhizoids. 5 5 5 5 5 laminae 1-stratose; costa percurrent to shortly excurrent, 5 5 Leaves straight, erect-spreading, 5 5 1 1 little changed when dry, smooth, /6– /4 the width of the leaves at base, rough with papillae 4–6 × 0.2–0.4 mm, most of the leaf or teeth on abaxial surface in distal half of leaf, abaxial tips deciduous and absent, ridges absent, with a row of guide cells, two weakly lanceolate, concave proximally, tubulose distally, apex developed stereid bands above and below, not extending to the leaf middle, adaxial and abaxial epidermal layers Dicranum · DICRANACEAE 419 of cells not differentiated or with a few cells enlarged in 26. Dicranum flagellare Hedwig, Sp. Musc. Frond., 130. both layers; cell walls between lamina cells not bulging; 1801 leaf cells smooth below, mammillose or usually abaxially Dicranum flagellare var. prorate or toothed in distal half of leaves, giving leaves a minutissimum Grout dull appearance; alar cells 1-stratose, differentiated or Plants in dense tufts, usually sometimes indistinctly differentiated, not extending to 5 bearing 2–6 dark green, stout, 5 costa; proximal laminal cells rectangular to oblong- 5 5 5 5 5 5 5 5 5 terete branchlets, 1–4 mm, in the 5 5 rectangular, not pitted or with few pits, (10–)20–32 5 55 5 5 5 5 5 55 5 5 5 5 5 555 5 5 5 axils of the distal leaves, each 5 5 5 5 (–50) × (4–)6–8(–13) µm; distal laminal cells usually 5 5 5 5 5 5 5 5 5 5 flagelliform branchlet with short, quadrate to short-rectangular, some cells transversely 5 elongate, not pitted, (5–)8–10(–26) × (2–)3–6(–9) µm. ovate to lingulate, broadly acute Sexual condition dioicous; male plants as large as females; to obtuse, ecostate or shortly and interior perichaetial leaves abruptly acuminate, indistinctly costate leaves closely appressed to the axis, convolute-sheathing. Seta 0.5–1.5 cm, solitary, rarely 2 yellowish green to dark green, glossy. Stems 0.5–6 cm, per perichaetium, yellowish to reddish brown. Capsule densely tomentose with light brown to reddish brown 1.2–2 mm, straight and erect to slightly inclined, rarely rhizoids. Leaves falcate-secund to nearly straight, crisped somewhat arcuate, smooth, striate when dry, light to weakly crisped when dry, smooth, (1–)2–4(–5) × 0.3– yellowish brown; operculum 1–1.5 mm. Spores 12–24 0.6 mm, concave below, tubulose above, from a µm. lanceolate or ovate-lanceolate base to an acuminate Capsules mature in spring. Rotting stumps and logs, subula, apex acute; margins smooth to serrulate above; tree bases (usually deciduous trees), occasionally soil or laminae 1-stratose; costa percurrent or sometimes ending 1 1 humus over rock, especially boulders and cliff ledges, in just before apex, /6– /4 the width of the leaves at base, dry to mesic woods, rarely swamps; 0–2800 m; B.C., smooth to ± rough with papillae or small teeth on abaxial Man., N.B., Nfld. and Labr. (Nfld.), N.W.T., N.S., surface near apex of leaf, abaxial ridges absent, with a Nunavut, Ont., P.E.I., Que., Sask.; Alaska, Ariz., Ark., row of guide cells, two stereid bands above and below Colo., D.C., Ill., Ind., Iowa, Ky., Maine, Md., Mass., not extending above the leaf middle, adaxial epidermal Mich., Minn., Mo., Nebr., N.H., N.Mex., N.Y., N.C., layer of cells not differentiated, the abaxial layer with Ohio, Okla., Pa., Tenn., Vt., Va., W.Va., Wis.; Europe; some cells differentiated; cell walls between lamina cells Asia. not bulging; leaf cells smooth or weakly abaxially prorate Dicranum montanum is one of the smallest species of above; alar cells 1-stratose, distinctly differentiated, not the genus in North America. It can best be recognized extending to costa; proximal laminal cells elongate- by the small plants, often only 1–2 cm, the dull, yellowish rectangular, not pitted or with few pits, (12–)24–38 green to dark green leaves that are strongly crisped when (–70) × (4–)10–15(–18) µm; distal laminal cells short- dry, the lanceolate, acuminate leaves that are subtubulose rectangular to quadrate, not pitted, (5–)12–14(–23) × to keeled above, rough on the abaxial surface near the (5–)7–12(–14) µm. Sexual condition dioicous; male apex, and the straight, erect capsules. Also aiding in its plants as large as females; interior perichaetial leaves recognition is the occasional presence of small, delicate, abruptly long-acuminate, convolute-sheathing. Seta 1– clustered branchlets with linear leaves that are readily 2.5 cm, solitary, yellowish to brown. Capsule 1.5–3 mm, detachable (probably a means of asexual reproduction), straight and erect or nearly so, striate when dry, yellowish which occur near the stem apices. The detached leaves brown to brown; operculum 0.5–l.6 mm. Spores 12–19 from the branchlets leave characteristic scars on the µm. denuded portions. Dicranum montanum can sometimes Capsules mature in spring. Rotting wood, especially be mistaken for D. flagellare, its closest relative, especially stumps and logs, base of trees, exposed tree roots, soil or when the latter species does not have its typical humus over boulders usually in mesic woods, sometimes flagelliform branchlets in the leaf axils or when they have swamps and bogs; 0–1300 m; Alta., B.C., Man., N.B., been overlooked. Dicranum montanum has leaves with Nfld. and Labr. (Nfld.), N.W.T., N.S., Ont., P.E.I., Que., a keeled subula in the distal half, with a strongly papillose Sask.; Ala., Ariz., Ark., Colo., Conn., Del., D.C., Fla., or toothed abaxial surface near the apex and distal leaf Ga., Ill., Ind., Iowa, Ky., La., Maine, Md., Mass., Mich., cells that are regularly quadrate. Dicranum flagellare, in Minn., Miss., Mo., Mont., Nebr., N.H., N.J., N.Y., N.C., contrast, has leaves with a tubulose subula above, with Ohio, Okla., Pa., R.I., S.C., S.Dak., Tenn., Vt., Va., W.Va., the abaxial surface usually smooth, and mostly short- Wis.; Mexico (Chihuahua, Coahuila, Durango, Jalisco, rectangular leaf cells. Nuevo León, Tamaulipas); West Indies; Central America (Costa Rica, Guatemala, Panama); Europe; Asia. Dicranum flagellare has been reported from Northwest Territories by W. L. Peterson (1979) and Arizona by 420 DICRANACEAE · Dicranum· Kiaeria

DICRANUM ° KIAERIA

I. M. Haring (1961). It is easily recognized because of the other types of asexual reproduction occurring in the the presence of unique axillary, flagelliform branchlets, genus. The only species likely to be confused with D. in clusters of 2–6, which some plants in each colony flagellare if the flagelliform branchlets are overlooked or almost always possess. The branchlets are deciduous and cannot be found is 25. D. montanum. For distinctions represent another type of asexual reproduction present see the discussion under that species. in species of Dicranum, e.g. deciduous branchlets with linear, crisped leaves in D. montanum, and deciduous Excluded Species: leaf tips in D. fragilifolium, D. tauricum and D. viride. C. E. Correns (1899) discussed the flagelliform branchlet Dicranum subporodictyon (Brotherus) C. H. Gao & T. type of asexual reproductive in D. flagellare as well as Cao (= Dicranodontium subporodictyon Brotherus)

12. KIAERIA I. Hagen, Kongel. Norske Vidensk. Selsk. Skr. (Trondheim) 1914: 109. 1915 · [For Frantz Caspar Kiaer, 1835–1893, Norwegian bryologist] Steven G. Newmaster

Dicranum sect. Falcata Bruch & Schimper, Bryol. Europ. 1: 117. 1847

Plants in erect, loose to dense tufts, green, yellowish or brownish, shiny or dull. Stems 1–2(–6) cm, erect-spreading, simple branches, sparsely radiculose near apex. Leaves lanceolate, gradually subulate, erect-spreading, sometimes falcate-secund; margins erect, entire, to serrulate near the tips; costa mostly excurrent, narrow, stereids poorly differentiated from median guide cells; distal Kiaeria · DICRANACEAE 421 laminal cells linear to subquadrate, smooth or slightly mammillose; basal laminal cells elongate, smooth, sometimes porose, alar cells usually brown and well differentiated. Sexual condition cladautoicous or gonioautoicous. Pericheatial leaves with a sheathing base. Seta solitary, 7–16 mm, erect, yellow turning red with age. Capsule suberect, curved, cylindric, smooth to furrowed when dry, indistinctly strumose; operculum obliquely rostrate; peristome single, of 16, teeth, divided halfway into two segments, papillose to vertically striolate or pitted striolate, red-brown. Calyptrae cucullate, occasionally rough at apex. Spores spheric, 14–24 µm, finely roughened, yellow to greenish yellow. Species ca. 6 (4 in the flora): North America, South America, Europe, Asia. A northern and alpine genus, Kiaeria occurs on siliceous rock or soil and is recognized by its medium-sized, Dicranum-like habit, with poorly differentiated stereid and guide cells. It may be confused with several other tufted acrocarps of similar size, such as Dicranoweisia, which has crisped, contorted leaves when dry; Oncophorus, which has distinctly strumose capsules and occurs on wood or humus; or Dicranella, which differs in its obliquely grooved capsules, and narrow hairlike leaves. The species in Kiaeria are difficult to distinguish, but some characters are useful for tentative field identification, including presence of grooves on capsule, leaf luster,

growth form, and habitat. ˆ ˆ

SELECTED REFERENCE Franklova, H. 2001. Distribution of the species of Kiaeria. Cas. Nár. Muz. Rada Prír. 170: 75–79.

1. Distal laminal cells elongate; capsule ribbed or grooved when dry. 2. Distal laminal cells not porose, basal laminal cells occasionally porose ...... 1. Kiaeria starkei 2. Distal and basal laminal cells porose ...... 4. Kiaeria glacialis 1. Distal laminal cells short, rectangular or quadrate; capsule smooth when dry. 3. Plants in loose tufts; leaves erect-spreading, flexuose ...... 2. Kiaeria blyttii 3. Plants in dense tufts; leaves falcate-secund ...... 3. Kiaeria falcata

1. Kiaeria starkei (Weber & D. Mohr) I. Hagen, Kongel. Kiaeria starkei occurs frequently on vertical rock Norske Vidensk. Selsk. Skr. (Trondheim) 1914: 114. surfaces, and is distinguished by the sessile male 1915 inflorescence, which is located close to the perichaetia. Dicranum starkei Weber & D. It can be separated in the field from the more terrestrial Mohr, Neues Bot. Taschenb. K. blyttii by its shiny, falcate leaves. Subalpine forms of 5 Anfänger Wiss. Apothekerkunst K. falcata are more commonly found on horizontal rock 1807: 189. 1807; Arctoa starkei surfaces and do not have grooved sporangia.

5 5 5 5 5 (Weber & D. Mohr) Loeske SELECTED REFERENCE Woolgrove, C. E. and S. J. Woodin. 1996. 5 5 5 5 5 5 Ecophysiology of a snow-bed bryophyte—Kiaeria starkei—during Plants in loose tufts, green to snowmelt and uptake of nitrate from meltwater. Canad. J. Bot. 74: yellow, mostly shiny. Stems 1–2 1095–1103. (–4) cm. Leaves mostly falcate- secund, curled at tips when dry, 2. Kiaeria blyttii (Bruch & Schimper) Brotherus, Laubm. lanceolate, gradually subulate, 2–4.5 mm, margins Fennoskand., 87. 1923 distally 1-stratose; costa 50–60 µm wide at base; distal Dicranum blyttii Bruch & Schimper, laminal cells mostly elongate (2–4:1), occasionally Kongl. Vetensk. Acad. Handl. 34: subquadrate, 7–9 µm wide, smooth or weakly 5 164. 1846; Arctoa blyttii (Bruch & mammillose-roughened; basal laminal cells elongate, 5 5 Schimper) Loeske; A. blyttii var. smooth, sometimes porose, alar cells strongly inflated 5 5 5 hispidula (R. S. Williams) Grout 5 5 5 5 and differentiated from the bordering small quadrate 5 5 5 5 5 Plants in loose or brown tufts, cells. Perichaetial leaves similar to the cauline. Perigonia 5 sessile, located just below perichaetia. Capsule distinctly dark green, dull. Stems 1–2 cm. ribbed when dry, urn 1.3–2 mm. Spores 13–21 µm. Branch leaves erect-spreading, Acid rock or sandy soil in rock crevices; subalpine to flexuose and somewhat crisped alpine elevations; Alta., B.C., Man., Nfld. and Labr. when dry, lanceolate, gradually subulate, 2–4 mm, (Nfld.), N.S., Que.; Alaska, Idaho, Mont., N.H., Oreg., margins occasionally distally 2-stratose; costa 35–50 µm Wash.; South America; Europe; Asia. at base; distal laminal cells mostly subquadrate (1–2:1), 422 DICRANACEAE · Kiaeria

KIAERIA ° ONCOPHORUS

8–11 µm wide, papillose on the dorsal surface; basal 3. Kiaeria falcata (Hedwig) I. Hagen, Kongel. Norske laminal cells elongate, smooth, sometimes porose, alar Vidensk. Selsk. Skr. (Trondheim) 1914: 112. 1915 cells gradually enlarged. Perichaetial leaves similar to Dicranum falcatum Hedwig, Sp. the cauline. Perigonia terminal on separate branches or Musc. Frond., 150. 1801; Arctoa located well below the perichaetia. Capsule not ribbed 5 falcata (Hedwig) Loeske

5 when dry, urn 1–1.2 mm. Spores 13–23 µm. 5 Plants in low, dense tufts, green, Soil in rock crevices or acid rock; alpine elevations; 5 5 5 5 dull. Stems 1–2(–4) cm. Leaves Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., N.S., Ont., 5 5 5 Que., Yukon; Alaska, Calif., Idaho, Maine, N.H., N.Y., mostly falcate-secund, lanceolate, Oreg., Wash.; Europe; Asia. gradually subulate, 2–3(–4 mm), Kiaeria blyttii is distinguished by position of the male margins distally 1-stratose; costa inflorescence at the ends of separate branches or well 50–60 µm wide at base; distal below the perichaetia, giving the appearance of separate laminal cells mostly rectangular (1–2:1), occasionally stems (polyoicous). In the field, it can be distinguished subquadrate, 7–9 µm wide, slightly papillose; basal from small sterile forms of K. starkei by its more terrestrial laminal cells elongate, smooth, sometimes porose, alar habit, spreading leaves, dull, dark green or brownish cells gradually differentiated. Perichaetial leaves similar color, and lack of grooves on the dry capsules. to the cauline. Perigonia sessile, located just below or occasionally distant from perichaetia. Capsule not ribbed when dry, urn 0.8–1.2 mm. Spores 12–19 µm. Acid rock or sandy soil; alpine elevations; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., N.S., Que., Yukon; Alaska, Oreg., Wash.; Europe; Asia. Kiaeria falcata is often found on horizontal rock surfaces in late-summer snowmelt areas. It grows in denser tufts than any other species of the genus. The epithet “falcata” refers to its falcate leaves, a character shared with K. starkei separating both species from Kiaeria· Oncophorus · DICRANACEAE 423

K. blyttii. It is distinguished from K. starkei by its shorter, µm wide, smooth or weakly mammillose-roughened; non-ribbed capsules and horizontal rather than vertical basal laminal cells elongate, porose, alar cells strongly rock habitat. inflated and differentiated. Perichaetial leaves similar to the cauline. Perigonia sessile, located just below 4. Kiaeria glacialis (Berggren) I. Hagen, Kongel. Norske perichaetia. Capsule distinctly ribbed when dry, urn 1.3– Vidensk. Selsk. Skr. (Trondheim) 1914: 125. 1915 2 mm. Spores 14–20 µm. Dicranum glaciale Berggren, Acta Soil over rock; mostly alpine elevations, occasionally subalpine; Greenland; Nfld. and Labr. (Nfld.), N.W.T., 5 Univ. Lund. 2(7): 19, figs. 1–9. Que., Yukon; Alaska; Europe. 5 1866; D. arcticum Schimper; 5 5 D. molle (Wilson) Lindberg Kiaeria glacialis is a large species that is found in late snowbeds and is similar to K. starkei, which is more 5 5 Plants robust, in loose tufts, slender and is commonly found on vertical rock surfaces. mostly green to yellow, shiny. A form of K. starkei occasionally occurs in late snowbeds, Stems 1–2(–4) cm. Leaves mostly but it has small, erect-spreading dark green leaves; K. falcate-secund, lanceolate, gradu- falcata also occurs in late snowbeds, but can be ally subulate, 2–4.5 mm, margins distinguished in the field by its short, non-grooved distally 1 or 2-stratose; costa 50–60 µm wide at base; capsules. distal laminal cells mostly elongate (2–5:1), porose, 7–9

13. ONCOPHORUS (Bridel) Bridel, Bryol. Univ. 1: 389. 1826 · [Greek onkos, tumor, and phoros, bearing, alluding to goiterlike swelling (struma) at base of capsule]

Steven G. Newmaster

Dicranum sect. Oncophorus Bridel, Muscol. Recent., suppl. 4: 53. 1819

Plants erect, in loose to dense tufts, green to yellow-brown, glossy to somewhat dull. Stems 1– 5(–8) cm, erect-spreading, simple or branching, radiculose proximally. Leaves lanceolate from an ovate base, gradually narrowed to abruptly subulate, crisped and curled when dry; margins plane or revolute, entire or often toothed at apex; costa strong, percurrent to slightly excurrent, narrow, with 2 stereid bands; distal laminal cells short-rectangular to subquadrate, smooth; basal laminal cells rectangular, elongate, smooth, alar cells poorly differentiated or sometimes slightly enlarged. Sexual condition autoicous; perichaetial leaves similar to the cauline; perigonia located below perichaetia. Seta 5–20(–22) mm, erect, twisted when dry, yellow to reddish yellow. Capsule inclined to horizontal, yellow-brown, arcuate, cylindric, urn 0.8–2 mm, constricted below the mouth, furrowed or smooth when dry, distinctively strumose; operculum rostrate, arcuate; peristome single, of 16 teeth divided halfway into two segments, vertically pitted-striolate proximally, papillose distally, red. Calyptra cucullate, smooth. Spores spheric, 14–25 µm, finely roughened, yellow to yellow-green. Species ca. 6 (3 in the flora): North America, Europe, Asia. Oncophorus is frequently encountered near stream habitats. The species may be confused with those of Dicranum, Dicranella, or Kiaeria, but are distinguished by distinctively strumose capsules and abruptly subulate leaves, which are strongly crisped-contorted when dry.

SELECTED REFERENCE Allen, B. H. 2000. The genus Oncophorus (Musci: Dicranaceae) in Maine. Evansia 17: 1–5.

1. Leaves keeled or folded, margins revolute, not sheathing at the base, gradually narrowed to the acumen ...... 2. Oncophorus virens 1. Leaves flat, margins plane, sheathing at the base, abruptly narrowed to the acumen. 2. Plants dark, not contorted when dry, seta usually less than 8 mm ...... 1. Oncophorus rauei 2. Plants lighter, contorted when dry, seta usually more than 8 mm ...... 3. Oncophorus wahlenbergii 424 DICRANACEAE · Oncophorus

1. Oncophorus rauei (Austin) Grout, Moss Fl. N. Amer. Ont., Que., Sask., Yukon; Ala., Alaska, Ariz., Ark., Calif., 1: 71. 1937 Colo., Conn., Del., D.C., Fla., Ga., Idaho, Ill., Ind., Iowa, Syrrhopodon rauei Austin, Bull. Kans., Ky., La., Md., Mass., Mich., Minn., Miss., Mo., Torrey Bot. Club 6: 74. 1876 Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Oreg., Pa., R.I., S.C., S.Dak., Tenn., Plants in loose tufts or extensive Tex., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Europe; mats, dark green to dark yellow- Asia. brown. Stems 1.5–3 cm. Branch A form of Oncophorus virens with coarsely toothed 5 leaves flexuose-spreading moist or 5 distal margins, was once treated as var. serratus and is 5 5 5 5 dry, ovate-lanceolate, abruptly often found on rocks in or near streams. The North subulate from ovate base, finely American material examined (as noted also by A. J. Grout serrate, 2.5–5 mm, subtubulose 1928–1940, vol. 2) varies considerably in the shape of above, sheathing at base, margins plane; costa stout, the leaf base, the extent of the revolute leaf margins, and mostly percurrent; laminal cells 2-stratose; distal laminal the presence or absence of alar cells in young plants. cells mostly subquadrate; basal laminal cells elongate, rectangular, incrassate, alar slightly differentiated. Seta 3. Oncophorus wahlenbergii Bridel, Bryol. Univ. 1: 400. 5–8 mm. Capsule yellow-brown, nearly smooth at 1826 maturity, 1–1.5 mm. Spores 16–19 µm. Capsules mature summer. Moist acidic rocks or cliffs Oncophorus wahlenbergii var. near streams or waterfalls; Ga., N.C., Pa., S.C., Tenn., 5 compactus (Bruch & Schimper) 5 Braithwaite 5 W. Va . 5 5 Oncophorus rauei, a rare species, has a south Plants soft, in small, loose tufts, 5 5 5 5 5 5 5 5 temperate distribution and is distinguished by its dark 5 5 5 light to very dark green. Stems 1– 5 5 5 5 5 55 5 5 5 5 5 5 5 55 5 5 5 5 5 5 5 5 color and rocky habitat. Although O. virens is also 5 5 5 55 2.5 cm. Branch leaves strongly 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 saxicolous, that species is distinguished by its folded 5 5 crisped and curled when dry, 5 leaves, which have revolute margins, and are gradually ovate-lanceolate, abruptly subulate, wihtout a sheathing base. Onchophorus subulate, entire or sometimes wahlenbergii is an epixylic species with leaves strongly serrated above, 3–6 mm, subtubulose above, strongly contorted when dry, the lamina is only 2-stratose at the sheathing at base, margins plane; costa mostly percurrent; margins, and the seta is usually longer. laminal cells mostly 1-stratose, 2-stratose only at margins; distal laminal quadrate to short-rectangular, incrassate; 2. Oncophorus virens (Hedwig) Bridel, Bryol. Univ. 1: basal laminal cells elongate, rectangular, incrassate, alar 399. 1826 cells not differentiated. Seta (8–)10–15(–21) mm. Capsule Dicranum virens Hedwig, Sp. Musc. pale yellow-brown, furrowed when dry, 0.8–2 mm, peristome bright red. Spores 14–20(–30) µm. 5 Frond., 142. 1801; Oncophorus

5 virens var. serratus (Bruch & Capsules mature late spring–early summer. Rotten 5 5 5 Schimper) Braithwaite logs along streams or in flooded areas, less commonly

5 5 on soil, humus, rock or bark at the base of coniferous 5 5 5 5 5 5 5 5 Plants in loose tufts, green to 5 5 5 5 trees (e.g., Thuja); Greenland; Alta., B.C., Man., N.B., 5 5 5 5 5 5 5 5 5 55 5 5 5 5 5 5 5 5 5 5 55 yellow-brown. Stems 1–5(–8) cm. 5 5 5 5 5 5 5 Nfld. and Labr. (Nfld.), N.W.T., N.S., Nunavut, Ont., 5 5 5 5 5 5 5 5 5 5 5 5 Leaves folded and sometimes 5 P.E.I., Que., Sask., Yukon; Ala., Alaska, Ariz., Ark., Calif., crisped when dry, flexuose when Colo., Conn., Del., D.C., Fla., Ga., Idaho, Ill., Ind., Iowa, moist, ovate-lanceolate, gradually Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., narrowed from ovate base, sometimes coarsely serrated Mo., Mont., Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., above, 2.5–5 mm, keeled distally, not sheathing at base, N.Dak., Ohio, Okla., Oreg., Pa., R.I., S.C., S.Dak., Tenn., margins revolute from base to leaf middle; costa stout, Tex., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Europe; mostly percurrent; laminal cells mostly 2-stratose; distal Asia. laminal cells mostly subquadrate; basal laminal cells A common epixylic species, Oncophorus wahlenbergii elongate, shorter at the margins, rectangular, incrassate, may be confused with the more saxicolous O. virens, alar cells enlarged and pellucid. Seta (7–)10–20(–22) mm. which has keeled, revolute, non-sheathing leaves that Capsule yellow-brown, slightly furrowed when dry, 1.5– gradually narrow to the subula. A compact, densely 2 mm. Spores 18–25 µm. tufted arctic-alpine form has been treated as var. Capsules mature late spring–early summer. Moist compactus and is often found on dry soil. rocks, rotten logs or soil; Greenland; Alta., B.C., Man., N.B., Nfld. and Labr. (Nfld.), N.W.T., N.S., Nunavut, Oreas· Paraleucobryum · DICRANACEAE 425 14. OREAS Bridel, Bryol. Univ. 1: 380. 1826 · [Greek Oread, mythological nymph of hills and mountains, alluding to alpine habitat]

William A. Weber

Plants in dense cushions, yellow-green distally, brown and compacted with red-brown radicles proximally. Stems forked. Leaves erect-spreading, crisped when dry, lance-acuminate, keeled, ending in a sharp, hyaline cell or short point; margins 2-stratose and narrowly revolute nearly to the apex, entire or somewhat irregular; costa prominent at base, smooth, ending near the leaf apex, entire to shortly excurrent; distal cells irregularly rounded-quadrate, smooth, thick- walled; basal cells narrowly rectangular, thick-walled, the alar cells not differentiated. Sexual condition autoicous; perigonial buds minute, at base of perichaetium; perichaetial leaves similar to stem leaves, ending in a short, hyaline awn. Seta straight or somewhat curved when dry, cygneous when moist, yellow. Capsule mostly erect when dry, pendent when moist, short-necked, symmetric, subglobose but becoming broader at the mouth and somewhat contracted at the middle when dry, orange-brown, strongly ribbed; annulus persistent; operculum obliquely rostrate from a convex base; peristome teeth inserted slightly below the mouth, lance-acuminate, brown at base, entire or occasionally somewhat perforated, rarely cleft, yellowish brown above, pale at the slender tips, vertically striolate throughout. Calyptra cucullate, smooth. Spores warty- papillose, brown. Species 1: North America, Europe, Asia. Oreas much resembles Cynodontium, but has a cygneous seta (when moist) and symmetric capsules. European specimens are more slender, with shorter leaves, but the differences are not significant.

SELECTED REFERENCES Steere, W. C. 1958. The discovery of Oreas martiana in arctic Alaska, a genus new to North America. Bryologist 61: 119–124. Weber, W. A. 1960. A second American record for Oreas martiana, from Colorado. Bryologist 63: 241–244.

1. Oreas martiana (Hooker) Bridel, Bryol. Univ. 1: 383. Capsules mature Jul–Aug. Alpine tundra, moderate 1826 to high elevations (to ca. 4200 m); Greenland; Alta., B.C.; Weissia martiana Hooker, Musci Alaska, Colo.; Europe; Asia. Oreas martiana is easily recognized by its golden color, 5 Exot. 2: plate 104. 1819

5 the very thick and deep polsters with only a few Stems 1.5–8.5 mm. Leaves 1.3–2 millimeters of living leaves and much brown material mm, distal laminal cells 6–10 µm. 5 5 below. In fruit it is unmistakable, with pendent, cygneous Perichaetial leaves 2–2.5 mm. Seta setae and strongly ribbed capsules. In the Front Range 3–7 mm. Capsule 0.6–0.9 mm; 5 of Colorado it appears to be particularly abundant. annulus of three rows of pale, thick-walled, quadrate cells; operculum 0.4–0.7 mm; peristome teeth 190–210 µm. Spores 16 µm.

15. PARALEUCOBRYUM (Lindberg ex Limpricht) Loeske, Allg. Bot. Z. Syst. 13: 162. 1907 · [Greek para-, near, and genus Leucobryum, alluding to resemblance] Robert R. Ireland Jr.

Dicranum subgen. Paraleucobrym Lindberg ex Limpricht in G. L. Rabenhorst et al., Krypt.-Fl. ed. 2, 4(1): 373. 1886 426 DICRANACEAE · Paraleucobryum

OREAS ° PARALEUCOBRYUM

Plants in loose to dense tufts, whitish green, grayish green or yellowish green, glossy. Stems 1– 4 cm, erect, simple or branched, rhizoids on stems below apex among leaves. Leaves subulate to narrowly lanceolate, gradually acuminate, subtubulose distally, erect-spreading to falcate- secund, especially at stem tips, little changed when dry, smooth; apices acute, sometimes deciduous; margins plane or incurved, serrate to serrulate distally to below leaf middle; laminae unistratose; costa single, percurrent, covering 1/2–2/3 of leaf base and all of subula, smooth or striate on abaxial surface, absent stereids, composed of 3–4 layers of large cells, hyaline, nonchlorophyllose cells intermingled with green, chlorophyllose cells, striations or ridges appearing as rows of teeth at high magnifications on abaxial surface of costa (P. longifolium); leaf cells pitted proximally, smooth, walls of medium thickness, median cells quadrate to rectangular, elongate proximally, alar cells inflated, extending to costa, 1-stratose, brown or sometimes hyaline. Specialized asexual propagation absent. Sexual condition dioicous; perigonia and perichaetia terminal; perigonial leaves ovate, concave, short-acuminate; perichaetial leaves with a broad base, abruptly narrowed to a long awn, inner ones sheathing. Seta solitary or rarely 2 per perichaetium, smooth, elongate, erect, straight to flexuose, twisted when dry, yellowish brown. Capsule erect to somewhat inclined, cylindric, straight, smooth, irregularly furrowed when dry, a few stomata at base, exothecial walls incrassate; annulus rudimentary; operculum rostrate, straight; peristome single, 16 lanceolate teeth, divided about halfway into 2 segments, papillose above, vertically to obliquely striate below, brown to reddish brown. Spores 22–33 µm, spherical to ellipsoidal, minutely papillose. Calyptra cucullate, smooth, naked, covering most of capsule, fugacious. Species 3 or 4 (2 in the flora): North America, Mexico, Europe, Asia. Paraleucobryum is a rather peculiar member of the Dicranaceae because of its unique leaf structure. In the leaf cross section there is an adaxial and abaxial layer of hyalocysts and a Paraleucobryum · DICRANACEAE 427 median layer of chlorocysts. Sometimes a few chlorocysts are in the abaxial layer of cells in some species. Brothera and Atractylocarpus have a very similar leaf structure and are the reason that the three genera are often placed in the same subfamily, Paraleucobryoideae. Both B. H. Allen (1999) and P. Müller and J.-P. Frahm (1987) presented extensive discussion on the relationship of Paraleucobryum to other members of the Dicranaceae. They also discuss what may be heterospory or bimodal spores in P. longifolium.

SELECTED REFERENCES Allen, B. H. 1999. The genus Paraleucobryum (Musci: Dicranaceae) in Maine. Evansia 16: 174–178. Barnes, C. 1958. The genus Paraleucobryum. Bryologist 61: 335–339. Ireland, R. R. 1994. Paraleucobryum. In: A. J. Sharp et al., eds. The moss flora of Mexico. Mem. New York Bot. Gard. 69: 147–148.

1. Costa with longitudinal striations, formed by rows of teeth evident at high magnifications, on abaxial surface (especially conspicuous in distal half); leaf margins serrulate near apex ...... 1. Paraleucobryum longifolium 1. Costa smooth on abaxial surface; leaf margins entire or rarely with a few teeth at apex ...... 2. Paraleucobryum enerve

1. Paraleucobryum longifolium (Ehrhart ex Hedwig) because of the falcate-secund leaves. The species was Loeske, Hedwigia 47: 171. 1908 reported from Alabama and Ohio by P. Müller and J.-P. Dicranum longifolium Ehrhart ex Frahm (1987). Paraleucobryum sauteri (Bruch & Schimper) Loeske 5 Hedwig, Sp. Musc. Frond., 130. has been considered a synonym by some bryologists (e.g., 5 1801; Campylopus canadensis 5 Kindberg; D. serratum Kindberg C. Barnes 1958; E. Lawton 1971). Gametophytically, it

5 is distinguished by the costa (R. S. Williams 1913, as 5 5 5 5 5 5 5 Plants whitish green to grayish 5 5 5 Dicranum sauteri Bruch & Schimper; P. Müller and J.-P. 5 55 5 5 5 5 5 5 5 green, sometimes yellowish, lighter 1 5 5 5 Frahm 1987) that is less than /3 the width of the leaf 5 5 5 5 5 in color at leaf bases. Stems 1– base compared to the costa that is more than 1/2 the width 4(–7) cm. Leaves spreading, of the leaf base in P. longifolium, which also means more usually falcate-secund, 4–8 × 0.2– rows of laminal cells in P. sauteri than in P. longifolium. 0.8 mm, margins usually serrulate in distal half; costa Müller and Frahm further distinguished P. sauteri by its 1 2 covering /2– /3 of leaf base, with longitudinal striations leaf cross section which has large median cells compared (ridges), appearing as rows of teeth at high to the smaller adaxial and abaxial layers of cells. magnifications, on abaxial surface, especially conspicuous Paraleucobryum longifolium, in contrast, according to in distal half, in cross section with adaxial hyalocysts, them, has small median cells in comparison to the larger median chlorocysts and abaxial hyalocysts with scattered adaxial and abaxial layers of cells. They also found that chlorocysts in some abaxial cells. Seta 8–20 mm. Capsule the peristome teeth of P. longifolium are divided only to 1.5–3 mm; operculum 1–2 mm. Spores 22–35 µm. the middle and inserted at the mouth, whereas in P. sauteri Capsules mature summer. Commonly on soil over the teeth are divided nearly to the base and inserted below noncalcareous boulders and cliffs, sometimes on tree the mouth of the capsule. Williams also used a peristome trunks, stumps, and rotten logs; moderate to high (400– difference to distinguish the two taxa. He found that in 2900 m); Greenland; Alta., B.C., Man., N.B., Nfld. and P. sauteri the peristome teeth are punctate or nearly Labr. (Nfld.), N.W.T., N.S., Ont., P.E.I., Que.; Ala., smooth on the exterior surface, whereas they are obliquely Alaska, Ariz., Colo., Maine, Mich., Minn., Mont., N.H., striate in P. longifolium. Müller and Frahm reported N.J., N.Mex., N.Y., N.C., Ohio, Pa., S.Dak., Tenn., Vt., specimens of P. sauteri only for western North America Va., W.Va., Wis.; Europe; Asia. through the Rocky Mountains from British Columbia to Paraleucobryum longifolium is best distinguished by Colorado, South Dakota, Arizona and New Mexico. I its 4–8 mm, whitish green to grayish green, glossy leaves have found the costa width in P. longifolium, however, that are falcate-secund, particularly at stem tips, slenderly to be quite variable in plants in eastern North America subulate with margins usually serrulate in distal half. The but some of the western North American plants do have costa has conspicuous fine striations or ridges, formed a narrow costa that fits the description of P. sauteri. I by small teeth when viewed at high magnifications, that also could not confirm the cross section difference are especially noticeable in the distal half even at low between the two taxa in the few North American magnifications with a dissecting microscope. The specimens that could be referred to P. sauteri. I have capsules are common, 1.5–3 mm, erect, cylindric, straight, decided not to recognize P. sauteri for this flora because smooth, with a 1–2 mm long-rostrate operculum. This I believe that a detailed study of the P. longifolium- species and the next somewhat resemble a Dicranum 428 DICRANACEAE · Paraleucobryum· Rhabdoweisia

P. sauteri complex is necessary, especially in regard to abaxial surface, in cross section with a row of adaxial the plants from the western part of the continent. hyalocysts, a row of median chlorocysts and a row of abaxial hyalocysts. Seta 10–20 mm. Capsule 2–3 mm; 2. Paraleucobryum enerve (Thedenius) Loeske, operculum 1–1.5 mm. Spores 15–20 µm. Hedwigia 47: 171. 1908 Capsules mature summer. Usually on soil or soil over Dicranum enerve Thedenius in C. J. boulders, noncalcareous outcrops and cliffs, sometimes in bogs and fens, rarely on stream banks; (150–4300 m); 5 Hartman, Handb. Skand. Fl. ed. 5, 5 393. 1849; D. albicans Greenland; Alta., B.C., N.W.T., Yukon; Alaska, Ariz., 5 5 Schwägrichen Colo., Mont., N.Mex., Utah, Wyo.; Mexico; Europe;

5 5 Asia. Plants whitish green to yellowish 5 Paraleucobryum enerve is best distinguished from P. 5 5 green. Stems 1–5 (–10) cm. 5 longifolium by its whitish green to yellowish green color, 5 5 Leaves erect-spreading to falcate- nearly smooth leaf margins that occasionally have a few secund, 2–8 × 0.5–1 mm, entire or teeth near the apex, and smooth costa. Sporophytes are with a few serrulations near apex; more rarely produced than they are by P. longifolium. costa covering ca. 9/10 or more of leaf base, smooth on

16. RHABDOWEISIA Bruch & Schimper, Bryol. Europ. 1: 97. 1846 · [Greek rhabdos, rod, presumably alluding to ribbed capsule, and genus Weissia, alluding to resemblance]

Wilfred B. Schofield

Plants in short loose tufts, perennial, dull green to brownish. Stems 0.2–3 cm, erect, forked by innovations, red-brown, without central strand, rhizoids smooth, at base of stem. Leaves oblong- lanceolate to narrowly lanceolate, obtuse to acute, keeled, strongly divergent when moist, crisped to sinuose when dry, 2–4 mm; costa subpercurrent, formed of rectangular cells; margins usually recurved, mainly serrate to serrulate along distal margins, with 1-celled irregular teeth; laminal cells smooth, proximal cells hyaline to brownish, rectangular, thin-walled, absent chlorophyll; distal cells, quadrate to irregular, usually broader than long, chlorophyllose except for the marginal teeth. Specialized asexual reproduction absent. Sexual condition autoicous; perigonial leaves short-elliptic; perichaetial leaves not differing from the vegetative. Seta solitary, erect, yellow, 2–6 mm, smooth. Capsule exserted, erect, brown to yellow-brown, 0.5–1 mm, symmetric, ovoid to oblong-cylindric, when dry and empty contracted below mouth and deeply 8-furrowed,; annulus absent; stomata at base of urn, phaneropore; operculum obliquely subulate, 0.5–1 mm; peristome single, 16 teeth joined by a low basal membrane, inserted slightly below 1 mouth, irregular and not forked, early deciduous. Calyptra cucullate, smooth, covering /2 of capsule. Spores nearly spherical to slightly angled, smooth, 16–20 µm, papillose. Species 6 (2 in the flora): North America, South America, Europe, Asia, Africa, Pacific Islands. Rhabdoweisia is known predominately from the Northern Hemisphere. A strong case has been made to include this, as well as other genera, in the family Rhabdoweisiaceae Limpricht.

SELECTED REFERENCE Lawton, E. 1961. A revision of the genus Rhabdoweisia. Bryologist 64: 140–156.

1. Leaves long-ligulate, 2–4 mm, broadly acute to obtuse, irregularly dentate in distal part, distal cells hexagonal, 14–20 µm wide, thin-walled relative to the lumina ...... 1. Rhabdoweisia crenulata 1. Leaves linear-lanceolate, 2–3.5 mm, gradually and narrowly acute, serrulate in distal part, distal cells mainly quadrate, often wider than long, 10–14 µm wide, somewhat incrassate ...... 2. Rhabdoweisia crispata Rhabdoweisia · DICRANACEAE 429

RHABDOWEISIA

1. Rhabdoweisia crenulata (Mitten) H. Jameson, Rev. 2. Rhabdoweisia crispata (Dickson ex Withering) Bryol. 17: 6. 1890 Lindberg, Acta. Soc. Sci. Fenn. 10: 22. 1872 Didymodon crenulatus Mitten, J. Bryum crispatum Dickson ex

Proc. Linn. Soc., Bot., suppl. 1: 23. 5 Withering, Syst. Arr. Brit. Pl. ed. 4,

1859 5 3: 816. 1801; Rhabdoweisia 5 5 Plants 0.5–3 cm. Leaves lingulate, denticulata (Bridel) Bruch, Schimper 5

5 5 5 & W. Gümbel 5 crisped when dry, 2–4 mm, 5 5 5 55 5 5 5 5 5 5 5 broadly acute to obtuse, margins 5 Plants 0.5–1 cm, occasionally 5 5 5 5 5 5 5 5 irregularly dentate distally, distal 5 5 taller. Leaves 2–3.5 mm, crisped cells 14–20 µm wide, irregularly when dry, narrowly lanceolate, hexagonal, rather thin-walled. gradually tapering to an acute Seta 3–4 mm. Capsule 0.6–1 mm; peristome teeth apex, margins serrulate with small teeth distally, distal narrowly lanceolate to subulate, smooth to faintly cells subquadrate, 10–14 µm wide, often wider than long, striolate, to 500 µm. Spores 18–20 µm. somewhat incrassate. Seta 2–4.5 mm. Capsule 0.7–1 Capsules mature summer. Wet, shaded ledge; mm; peristome teeth linear to linear-lanceolate, moderate elevations (ca. 1100 m); of conservation broadened at base, smooth to striolate, perforate or concern; N.C.; Europe; Asia; Pacific Islands (Hawaii). irregular. Spores 15–21 µm. On the North American continent, this species is Capsules mature summer. Mainly on dampish, shaded known from a single locality in the Appalachian cliffs; 1–2500 m; Greenland; B.C., Man., Nfld. and Labr. Mountains (R. H. Zander 1966). (Nfld.), N.W.T., N.S., Ont., Que., Yukon; Ala., Alaska, Ark., Colo., Conn., Ga., Ill., Ky., Maine, Mass., Mich., Minn., Mo., N.H., N.J., N.Y., N.C., Pa., Tenn., Vt., Va., Wis.; South America (Bolivia); Europe; Asia; Pacific Islands (Hawaii, Juan Fernández Islands). In eastern North America, Rhabdoweisia crispata exhibits a wide distribution from the coast to more 430 DICRANACEAE · Rhabdoweisia· Symblepharis continental areas, but on the Pacific coast it is primarily subquadrate and broader than long while those of R. in near-coastal oceanic climates. It is distinguished from crenulata are slowly rehydrated, remaining somewhat R. crenulata mainly on the basis of leaf shape: narrowly collapsed until hydrated, and are quadrate to hexagonal. lanceolate and gradually acute in the former, lingulate The leaves of R. crispata are serrulate with small teeth, and broadly acute to obtuse in the latter. The distal leaf while R. crenulata has dentate leaves with larger, widely cells of R. crispata are rapidly rehydrated, and generally spaced teeth.

17. SYMBLEPHARIS Montagne, Ann. Sci. Nat., Bot., sér 2, 8: 252. 1837 · [Greek sym-, united, and blepharis, eyelash, alluding to peristome teeth each parted in distal half into two ciliate divisions]

Patricia M. Eckel

Plants moderate in size, in dense, yellow-green to golden yellow shining tufts. Stems 2–4.5 cm, erect, simple or forked, densely foliate, rhizoids smooth, red-brown, sparse, concentrated along the proximal portion of the stem, central strand present. Leaves crisped-contorted when dry, widely spreading-flexuose when wet, subfalcate, abruptly narrowed from a flaring, obovate, strongly shouldered, sheathing base into a long, linear-lanceolate, canaliculate subula; apex subulate, not deciduous, distal lamina serrate abaxially; margins canaliculate, entire at the base, becoming distantly low-denticulate distally and strongly serrate near apex, 1- or irregularly 2- stratose; costa single, narrow, filling less than 1/3 of the leaf base, percurrent to shortly excurrent, smooth but sharply toothed abaxially near the apex, adaxial epidermis absent, stereid bands 2, adaxial band disappearing in apex; laminal cells plane, unpitted, smooth, differentiated, basal cells long-rectangular, thin to vesciculose, not differentiated along costa or basal angles, distal cells subquadrate, thick-walled. Specialized asexual reproduction none. Sexual condition autoicous; antheridia gemmiform, occurring at various distances below the perichaetium; perichaetial leaves similar to apical stem leaves but with higher sheathing bases. Seta 1–4 per perichaetium, erect, ca. 1 cm, slender, smooth, yellow. Capsule erect, symmetric, slenderly cylindric, straight or slightly arcuate, smooth or irregularly striate when dry, base gradually tapering to the seta, not strumose, annulus indistinctly differentiated; operculum obliquely long- rostrate; peristome teeth deeply inserted, 16, in 8 groups each composed of 2 teeth joined at the base and each tooth distally divided to above the middle into two rami, the 2 inner, neighboring rami shorter and narrower than the outer rami, vertically striolate at base and 1 strongly papillose in the upper /3; deep red. Calyptra narrow, cucullate, rough in the apex, 1 entire at the base, naked, covering the distal /2 of the capsule, fugacious. Spores 12–18 µm, smooth to very finely papillose, irregularly ovoid. Species 12 (1 in the flora): North America, Mexico, Central America, South America, Asia, Africa. Nine species of Symblepharis occur in southeastern Asia, two in the Neotropics, and one in tropical Africa. The genus is recognized by wide-spreading leaf tips that are crisped when dry, the linear-lanceolate lamina abruptly distinct from a flaring obovate-sheathing base, and, when they do occur, setae clustered. The distinctive striate proximal and papillose distal portions of the peristome teeth, and the grouping of the teeth into eight tooth-pairs are also characteristic. The organization of the peristome is somewhat more complex than descriptions usually indicate, the eight groups of paired teeth being separated by “a space equivalent to one tooth between each pair” (H. C. Gangulee 1969+, fasc. 2). The gametophyte of the genus is nearly identical with that of the common and widespread Oncophorus wahlenbergii. Since both species display bistratose leaf margins, this is not a good character to differentiate them (see discussion below). Symblepharis · DICRANACEAE 431

SYMBLEPHARIS ° BRUCHIA

In S. vaginata the capsule is erect, long-cylindric, symmetric, and tapering gradually to the seta, and is essentially smooth wet or dry. The capsule of O. wahlenbergii is inclined to horizontal, short-ovoid, asymmetric, and with a goiterlike swelling at the base (as is the case for all species of Oncophorus), and is generally distinctly furrowed when dry.

SELECTED REFERENCES Ireland, R. R. 1994b. Symblepharis. In: A. J. Sharp et al., eds. The moss flora of Mexico. Mem. New York Bot. Gard. 69: 156. Salmon, E. S. 1898. A revision of the genus Symblepharis, Montagne. J. Linn. Soc., Bot. 33: 486–501.

1. Symblepharis vaginata (Hooker) Wijk & Margadant, subquadrate, 10–12.5 × 6–8 µm, thick-walled, 1-stratose Taxon 8: 75. 1959 except for irregular 2-stratose patches on the distal Didymodon vaginatus Hooker, margins. Seta 10–12 mm, straight, single or occasionally Icon. Pl. 1: plate 18, fig. 4. 1837; clustered in groups of 2–4. Capsule yellow, 3–4 mm, Symblepharis helicophylla Montagne deep red at the mouth, smooth to weakly and irregularly striate in older capsules; peristome inserted below the Stems ca. 0.8–1.2(–3) cm, leaves mouth; operculum ca. 0.8 mm. spreading, flexuose, and sub- Capsules mature winter. Rotting punky, sometimes falcate when wet, crisped and charred logs, less commonly on bark at base of trees, 5 5 tortuose when dry, sparingly rock, soil in coniferous forests; high elevations (2200– radiculose at the base. Leaves 6– 3100) m, occasionally to 3800 m; Ariz., N.Mex.; Mexico; 7 mm, leaf base broader at the Central America (Costa Rica, Guatemala, Panama); Asia undulating shoulders than the extreme base, with a (China, India, Japan, Korea, Papua New Guinea). narrow, fragile decurrency, margins plane, distantly Stem growth beyond an apparently lateral sporophyte denticulate distally, sharply serrulate near the apex; basal is actually the extension of a subperichaetial branch cells smooth, elongate-rectangular to linear, 65–85 µm, (sympodial branching). As in Symblepharis, all three narrowly long-rectangular, thin-walled, not porose, clear, species of Oncophorus in North America have bistratose medial lamina cells irregularly short-rectangular to leaf margins; the blade of O. rauei is bistratose 432 DICRANACEAE · Symblepharis throughout its distal lamina in addition to the margins. to distinguish these two species when sterile is the leaf Oncophorus virens does not have a clasping leaf base, cross section. The leaf of S. vaginata is histologically and the distal portion of the blade tapers gradually to reduced: the adaxial epidermis is absent at mid leaf and, the apex. The gametophyte of S. vaginata is, however, distally even the adaxial stereid band vanishes, leaving nearly identical with that of O. wahlenbergii, except it is only the guide cell layer. In O. wahlenbergii the mid leaf of a relatively clear yellow, whereas O. wahlenbergii is a costa is robust, and the adaxial epidermis layer is sooty, fuscous yellow. One useful character with which prominent.