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Flora of North America, Volume 27, 2007

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Flora of North America, Volume 27, 2007 254 GRIMMIACEAE · Grimmia weakly striate, exothecial cells thin-walled, annulus basic rock, has setae 2–3 mm, shiny, spherical capsules, present, operculum rostrate, peristome teeth orange, fully- and a mammilate operculum. Only the most proximal developed to irregularly cleft at apex, papillose. Calyptra stem leaves of G. orbicularis are muticous and the basal mitrate. juxtacostal cells have thick, nodulose walls. Dry acidic to basic rock; low to moderate elevations (60–1000 m); B.C.; Calif., Oreg., Wash.; Mexico; Eurasia; 36. Grimmia muehlenbeckii Schimper, Syn. Musc. Eur., n Africa; Pacific Islands (Hawaii). 212. 1860 Grimmia lisae is a thermophilous species with a Grimmia hermannii H. A. Crum; G. preference for subtropical coastal areas. In North trichophylla var. tenuis (Wahlenberg) America, it occurs along the west coast, from Vancouver Wijk & Margadant Island south to Mexico. From that region, it has been Plants in blackish green tufts. described frequently as a new species. It is closely related 5 5 5 5 Stems 1–2.5 cm, central strand 5 to G. trichophylla, but is distinguished by somewhat 5 5 5 5 5 5 5 shorter and broader leaves that are straight and appressed present. Leaves loosely appressed, when dry and recurved to squarrose when moist, and by twisted when dry, erectopatent a reniform costa. Furthermore, it is characterized by a when moist, ovate-lanceolate, grass-green mid leaf areolation with small, rounded, tapering to acute apex, 2–3 × 0.6– frequently oblate cells with straight walls. 0.8 mm, keeled, margins recurved in mid leaf on both sides, awns short, denticulate, in perichaetial leaves stout 35. Grimmia moxleyi R. S. Williams in J. M. Holzinger, and often decurrent, costa channeled distally, projecting Musci Acroc. Bor.-Amer., 600. 1926 at abaxial side, angled to bluntly winged; basal juxtacostal laminal cells short- to long-rectangular, yellowish, Plants in frequently extended nodulose, thick-walled; basal marginal laminal cells short- mats, blackish green. Stems 1–1.5 rectangular with thickened transverse walls; medial mm high, small central strand laminal cells quadrate to short-rectangular, sinuose, thick- present. Leaves erect with slightly walled; distal laminal cells 1-stratose with 2-stratose incurved tips when dry, erect- ridges, margins 2-stratose. Gemmae rare, in clusters, spreading when moist, oblong, 5 5 short-stalked, in distal leaf axils. Sexual condition 5 broadly rounded and muticous at dioicous. Seta arcuate, 2–3 mm. Capsule occasionally apex, 1.5–2 × 0.4–0.6 mm, keeled, present, exserted, globose, shiny, brown, smooth to margins plane to recurved, awns slightly striate, exothecial cells thin-walled, annulus absent, only present in perichaetial leaves, costa weak present, operculum rostrate, peristome teeth purple, fully- proximally, projecting at abaxial side, perichaetial leaves developed or slightly split distally, papillose. Calyptra longer and with awns; basal juxacostal laminal cells short- mitrate. to long-rectangular, straight, thin-walled; basal marginal Shaded acidic rock, often along lakes; moderate to laminal cells short- to long-rectangular, thin-walled; high elevations (200–2000 m); B.C., Nfld. and Labr. medial laminal cells rounded-quadrate, ± sinuose, thin- (Nfld.), Ont.; Idaho, Maine, Mass., Mich., Minn., N.H., or thick-walled; distal laminal cells 2-stratose. Gemmae Oreg., Wash., Wis.; Europe; e Asia (Japan); Africa (South absent. Sexual condition autoicous. Seta flexuose to Africa). arcuate, 1–1.5 mm. Capsule usually present, exserted, Grimmia muehlenbeckii is closely related to G. chestnut brown, oblong-ovoid, wrinkled-plicate when trichophylla, and in the past it was frequently regarded dry, exothecial cells thin- to thick-walled, annulus present, as a subspecies or variety of that taxon. H. Deguchi operculum conical, peristome teeth yellowish, split and (1978) treated it as a distinct species, followed by A. J. E. perforated, papillose. Calyptra cucullate. Smith (1992). Greven agrees with those treatments. He Dry acidic rock; moderate elevations (500–1500 m); has seen many specimens that are remarkably uniform of conservation concern; Ariz., Calif., Nev.; Mexico. and easy to distinguish from G. trichophylla by their Grimmia moxleyi is endemic to the southwestern small, globose, shiny, dark brown capsules with purple, United States and northwestern Mexico. A entire peristome teeth. In contrast, the capsules in G. thermophilous species from acidic rock, it is autoicous trichophylla are oblong-ovoid, larger and longer than in and usually richly provided with capsules. The species is G. muehlenbeckii, dull, yellowish brown, and the characterized by growing in flat, easily disintegrating peristome teeth are orange and cleft. The gametophyte patches with short, keeled, muticous stem leaves that differs from that of G. trichophylla by its blackish green contrast with its much larger awned perichaetial leaves. tufts, angled costa with blunt wings protruding on the J. Muñoz (2000) synonymized G. moxleyi with G. abaxial side, and ovate-lanceolate leaves with stout, orbicularis. Although there are some similarities, G. denticulate awns that are often decurrent. orbicularis is much larger, grows in dense cushions on Grimmia · GRIMMIACEAE 255 37. Grimmia orbicularis Bruch in J. E. Smith et al., Engl. 38. Grimmia pulvinata (Hedwig) Smith in J. E. Smith et Bot., suppl. 4: plate 2888. 1844 al., Engl. Bot. 24: plate 1728. 1807 Plants in hoary, usually Fissidens pulvinatus Hedwig, Sp. hemispherical cushions, grayish Musc. Frond., 158, plate 40, figs. green. Stem 2–5 cm, central strand 1–3. 1801; Grimmia decipiens var. present. Leaves appressed and hendersonii (Renald & Cardot) 5 twisted when dry, erect when 5 Sayre; G. indianensis (Sayre) H. A. 5 5 5 5 5 5 moist, broadly lanceolate, 5 5 Crum; G. pulvinata var. africana 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 abruptly contracted into awn, 2– 5 Hooker f. & Wilson; G. subcurvula 5 2.5 × 0.4–0.6 mm, keeled, margins 5 Kindberg; G. trichophylla var. recurved in the middle of the leaf indianensis Sayre on one or both sides, awns short to long, smooth to Plants in hemispherical hoary cushions, grayish green. denticulate, costa weak proximally, projecting on abaxial Stems 1–3 cm, central strand present. Leaves flexuose side; basal juxtacostal laminal cells short- to long- when dry, erect when moist, lanceolate, 1–1.7 × 0.3–0.6 rectangular, sinuose-nodulose, thick-walled; basal mm, keeled, margin recurved on both sides nearly from marginal laminal cells short- to long-rectangular, straight, base to apex, rarely only one margin recurved, abruptly with thickened transverse walls; medial laminal cells contracted into short to long, smooth to denticulate awn, subquadrate, sinuose, thick-walled; distal laminal cells costa weak proximally, projecting on abaxial side; basal 1-stratose. Gemmae absent. Sexual condition autoicous. juxtacostal laminal cells quadrate to short-rectangular, Seta arcuate, 2–3 mm. Capsule usually present, exserted, thin-walled; basal marginal laminal cells quadrate to bent down into the cushions by the arcuate setae, short-rectangular, thin- or thick-walled; medial laminal yellowish brown to chestnut brown, globose to ovoid, cells rounded-quadrate, slightly sinuose, thin-walled; smooth to weakly ribbed, when dry and empty wide- distal laminal cells 1-stratose, margins 2-stratose. mouthed, exothecial cells thin-walled, annulus present, Gemmae absent. Sexual condition autoicous. Seta operculum mammillate, peristome teeth orange, broad, arcuate, 3–4 mm. Capsule usually present, exserted, cribrose and irregularly cleft at apex. Calyptra cucullate. ellipsoid to obloid, brownish and ribbed when empty and Dry basic rocky substrates such as limestone, basalt, dry, exothecial cells thin-walled, annulus present, and mortar; moderate to high elevations (200–2000 m); operculum rostrate, peristome teeth reddish, fully- Ariz., Calif., Mont., Nev., Utah; Mexico; Central America developed, papillose. Calyptra mitrate. (Guatemala); South America (Argentina, Chile); Europe; Various substrates, from acidic to basic rock, old n Africa; Pacific Islands (New Zealand); Australia. mortar, tree trunks; low to high elevations (0–3000 m); Grimmia orbicularis is a thermophilous species with Alta., B.C., Ont.; Ariz, Calif., Colo., Idaho, Ind., Iowa, a preference for sunny, basic substrates. In North America Kans., Md., Mich., Miss., Mo., Mont., Nev., N.Mex., it is known only from scattered localities in the Southwest. Okla., Oreg., S.Dak., Tex., Utah, Wash., Wyo.; Mexico; It may be confused with G. pulvinata, as both species South America (Argentina, Chile, Uruguay); Eurasia; form comparable hemispherical cushions and usually Africa; Pacific Islands (New Zealand); Australia. grow in the same habitat. However, they differ markedly Grimmia pulvinata is the most common species of the in both gametophytic and sporophytic characters. genus. It has a nearly cosmopolitan distribution, and is Grimmia orbicularis has leaves with short- to long- a pioneer on various substrates, even on the trunks of rectangular basal juxtacostal cells with thick and trees. However, in the eastern part of North America, it nodulose lateral walls, and 1-stratose margins while G. occurs only in a few scattered localities (H. A. Crum pulvinata has leaves with quadrate to short-rectangular, 1977). It is easily recognized by its neat hemispherical thin-walled, basal juxtacostal cells and 2-stratose margins. cushions with abundant capsules. It may be confused Grimmia orbicularis has globose
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