254 · 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 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 to ovoid capsules with with G. orbicularis, which grows in similar habitats and broad, orange, cribrose and cleft peristome teeth, and is known to co-occur with G. pulvinata. However, as mammillate opercula while G. pulvinata has obovoid discussed under G. orbicularis, it differs in gametophytic capsules with fully developed dark red peristome teeth as well as sporophytic characters. Some forms with more and rostrate opercula. acuminate leaf tips may be confused with G. trichophylla, but the thin-walled, short-rectangular basal cells and the small, rounded mid leaf cells distinguish G. pulvinata. 256 GRIMMIACEAE · Grimmia

39. Grimmia ramondii (Lamarck & de Candolle) like areolation, and the distinct costal wings distinguish Margadant, Lindbergia 1: 128. 1972 it easily from those species. Pterigynandrum ramondii Lamarck 40. Grimmia torquata Drummond, Musc. Scot. 2: no. 5 & de Candolle, Fl. Franç. ed. 3, 2: 28. 1825 5 462. 1805; Dryptodon patens 5 (Hedwig) Bridel; Grimmia curvata Grimmia pellucida Kindberg;

5 5 5 (Bridel) De Sloover, G. serrata 5 G. prolifera Müller Hal. &

5 5 5 5 5 5 Kindberg; Racomitrium patens Kindberg; G. pseudotorquata 5 5 (Hedwig) Huebener 5 Kindberg; G. tortifolia Kindberg

5 5 5 5 5 5 Plants in robust, loose, arched- 5 Plants in soft, readily disinte- 5 5 5 5 5 ascending, readily disintegrating 5 grating cushions, yellow-green to 5 patches, brownish green distally, blackish proximally. 5 brownish, occasionally light green, Stems 5–10 cm, central strand absent. Leaves loosely blackish to red-brown inside. appressed when dry, patent when moist, broadly ovate- Stems 1–4 cm, small central strand lanceolate, tapering to an acute, slightly toothed apex, present. Leaves contorted when dry, patent when moist, 2–3 × 0.4–0.6 mm, keeled, margins recurved on both lanceolate, 1.5–2 × 0.3–0.5 mm, keeled, margins slightly sides, awns absent, costa stout, with two abaxial lamellae recurved proximally, plane distally, awns very short and (or wings) forming parallel ridges along its length; basal smooth, occasionally absent, costa weak proximally, juxtacostal laminal cells yellow, linear, sinuose, thick- projecting on abaxial side; basal juxtacostal laminal cells walled; basal marginal laminal cells quadrate to short- linear, extremely sinuose, thick-walled; basal marginal rectangular, slightly thick-walled; medial laminal cells laminal cells long rectangular, thin-walled, hyaline; medial quadrate to rectangular, sinuous, thick-walled; distal laminal cells rectangular, extremely sinuose, thick-walled; laminal cells 1-stratose, margins 2-stratose. Gemmae distal laminal cells 1-stratose. Gemmae brown, absent. Sexual condition dioicous. Seta cygneous to multicellular, present on the abaxial side of distal leaves. arcuate at maturity, flexuose when old, 3–5 mm. Capsule Sexual condition dioicous. Seta slightly curved, straight occasionally present, exserted, obloid, yellowish green when dry, 3–5 mm. Capsule sporadically present, to yellowish brown, plicate when empty, exothecial cells exserted, ovoid, brown, smooth, striate when dry and thin-walled, annulus present, operculum rostrate, empty, exothecial cells thin-walled, annulus present, peristome teeth purple, divided nearly to base into two operculum rostrate, peristome teeth yellowish, short, split partly adhering segments, papillose. Calyptra mitrate. in distal part, weakly papillose. Calyptra mitrate. Dry to damp acidic rock; low to high elevations (0– Damp, frequently vertical faces of acidic rock; 2000 m); Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), moderate to high elevations (200–4000 m); Greenland; N.W.T.; Alaska, Calif., Idaho, Mich., Mont., Oreg., Alta., B.C., Nfld. and Labr., N.W.T., Ont., Que., Yukon; Wash.; Europe; Asia (China, Japan); n Africa. Alaska, Ariz., Calif., Colo., Idaho, Mont., Nev., Oreg., Grimmia ramondii occurs near sea level in arctic areas Wash., Wyo.; Mexico; Eurasia; n Africa; Pacific Islands and along the West Coast. In the latter region, however, (Hawaii). it is more common in mid-elevation montane sites and Grimmia torquata is a montane-alpine species with a may also be found above the tree line. It differs from preference for shaded habitats. It grows in hemispherical most Grimmia species by its lack of awns, winged costa, cushions on steep, damp rock walls. Preferred substrates and tall, loose growth form. At first sight, G. ramondii are granite, gneiss, quartzite, and schist. Its distribution has the habit of a Racomitrium. However, the sporophyte reaches from near sea level in the Arctic to above 4000 has an arcuate rather than straight seta, and the capsules m on Mexican and Hawaiian volcanoes. The species is are striate rather than smooth. Because of its habit and easily recognized by its yellow-brown cushions of plants, the absence of a central strand, K. G. Limpricht which have contorted leaves when dry, and their brown ([1885–]1890–1903, vol. 1) placed it, together with G. gemmae, which are borne at the bases of older leaves. In hartmanii and G. atrata, in Dryptodon, a genus densely shaded habitats, the cushions are light green; on intermediate between Grimmia and Racomitrium. There exposed rock, they are usually brown. Although the are many other Grimmia species, however, without a species has a wide distribution, it is seldom present in central strand, and G. hartmanii and G. atrata have quantity, usually growing in a small number of cushions already been removed from Dryptodon, so there is no in one habitat. Sporophytes are very rare worldwide and reason to maintain that monotypic genus. Grimmia have not been found in eastern North America (H. A. ramondii may be confused with G. hartmanii and G. Crum and L. E. Anderson 1981). elatior. However, the lack of awns, the Racomitrioideae- Grimmia · GRIMMIACEAE 257

41. Grimmia trichophylla Greville, Fl. Edin., 235. 1824 Mont., Nev., Okla., Oreg., S.Dak., Utah, Vt., Wash., Wyo; Plants in dense to loose patches, Mexico; Eurasia; Pacific Islands (Hawaii); Australia. yellowish green to dark green. In North America, Grimmia trichophylla is principally Stems 2–4 cm, central strand a lowland species, occurring in the mountains up to about present. Leaves loosely appressed, 1000 m., rarely higher. In the Southern Hemisphere, it 5 slightly twisted when dry, erecto- may be found up to 4000 m. In New Zealand, the species 5 5 5 5 5 is common, and in contrast to G. trichophylla in North 5 5 5 patent when moist, lanceolate to 5 5 5 5 5 America, frequently bears capsules. The New Zealand 5 5 oblong-lanceolate, tapering to acute apex, 2–3.5 × 0.3–0.4 mm, plants are usually smaller than American specimens, and usually sharply keeled, margins the leaves are frequently contorted. The nearly recurved on one or both sides, plane to erect distally, cosmopolitan G. trichophylla has many phenotypes, and awns variable, short to long, smooth to denticulate, not numerous subspecies and varieties have been described. conspicuously flattened at base, costa firm, projecting In damp and shaded habitats, the awns may be short, on abaxial side; basal juxtacostal laminal cells long- just as in dry unfavorable habitats at high altitudes, where rectangular (rarely short-rectangular), ± nodulose, thick- stunted specimens may occur with small, short leaves and walled; basal marginal laminal cells short- to long- reduced awns, or even with muticous leaves. Grimmia rectangular, with thickened transverse walls; medial trichophylla has frequently been confused with related laminal cells quadrate to short-rectangular, slightly species such as 36. G. muehlenbeckii and 34. G. lisae sinuose, thick-walled; distal laminal cells 1-stratose, (see discussions thereunder for identification details). occasionally with 2-stratose ridges. Gemmae clusters Robust forms of G. trichophylla have been mistaken for occasionally present in distal leaf axils. Sexual condition G. austrofunalis (H. C. Greven 1997, 2003), which does dioicous. Seta arcuate, 2–4 mm. Capsule occasionally not occur in North America. Although some of those present, exserted, oblong-ovoid, yellowish green to plants have leaves of equal length along the stem, stramineous, striate when dry, exothecial cells thin-walled, characteristic of G. austrofunalis, they also have both annulus present, operculum rostrate, peristome teeth leaf margins recurved, and the medial and outer basal yellowish, papillose, deeply split and perforated. Calyptra laminal cells are longer and more robust than in that mitrate. species. Dry, acidic rock; moderate to high elevations (200– 2000 m); B.C.; Ariz., Calif., Colo., Idaho, Maine, Mo.,

2e. GRIMMIA (subg. undetermined)

42. Grimmia mollis Bruch & Schimper, Bryol. Europ. 3: teeth purple, perforated, fully-developed, papillose. 133, plate 253. 1849 Calyptra cucullate. Hydrogrimmia mollis (Bruch & Wet acidic rocks in alpine and boreal habitats; moderate to high elevations (1000–4100 m); Greenland; 5 Schimper) I. Hagen; Grimmia 5 evansii E. Britton Alta., B.C., Que., Yukon; Alaska, Calif., Colo., Mont., 5 Wash.; Eurasia. Plants in loose patches, green. 5 5 Grimmia mollis, discovered by Schimper in the Tirol 5 5 Stems 1–5 cm, central strand 5 of Austria, is a characteristic arctic-alpine species,

5 present. Leaves oblong to ovate- 5 frequently growing in glacial streams, sometimes over lanceolate, 2–3 × 0.8–1 mm, reaches of hundreds of meters. Although the species is margins plane, incurved distally, widespread in the Northern Hemisphere, it is nowhere muticous, concave, apex rounded common. It occurs exclusively above the treeline. Because to cucullate, costa not projecting on abaxial side, of its unique subpercurrent costa, uniform lamina of subpercurrent; basal juxtacostal laminal cells quadrate, quadrate, thin-walled cells, and peculiar ecological niche, thin-walled; basal marginal laminal cells quadrate, thin- it cannot easily be confused with any other species. walled; medial laminal cells quadrate, thin-walled; I. Hagen (1909) placed it in a monotypic genus, juxtacostal and marginal distal laminal cells 1-stratose. Hydrogrimmia. This concept was followed by M. F. V. Sexual condition dioicous. Seta straight, 2–3 mm, Corley et al. (1981). However, there are no significant yellowish. Capsule sporadically present, exserted, morphological characters to separate it from Grimmia, yellowish, ovoid to oblong, smooth, exothecial cells thick- prompting H. C. Greven (2003) to follow, in agreement walled, annulus absent, operculum rostellate, peristome with H. A. Crum and L. E. Anderson (1981), Bruch and 258 GRIMMIACEAE · Grimmia· Coscinodon

Schimper’s original concept. This dioicous species grows Volcanic rock; high elevations (3500 m); of in separate male and female cushions, and capsules are conservation concern; Calif. produced very rarely. Grimmia shastae is known only from Mt. Shasta. Its discovery demonstrates that the bryoflora of California 43. Grimmia shastae Greven, Grimmias, 208. 2003 (as is still not fully known. It also shows that the state is shastai) exceptionally rich in endemic species of Grimmia: G. Plants in hoary tufts, brownish mariniana, G. nevadensis, G. serrana, and G. shastae. green. Stems 0.3–1 cm. Leaves Because of its abundantly present gemmae on the adaxial ovate to broadly oblong- surface of the distal lamina, the latter is not likely to be lanceolate, 1–1.5 × 0.5–0.6 mm, confused with any other species of the genus. While larger towards stem tips, not gemmae are not rare in Grimmia, they are usually keeled, frequently plicate in produced on leaf tips (G. anomala, G. hartmanii), on 5 extreme apex, margins plane, the abaxial side of the leaf (G. torquata, G. trichophylla), incurved distally, long-awned, or in leaf axils (G. trichophylla, G. muehlenbeckii). In decurrent, costa weak proximally, no other Grimmia species are gemmae produced in broadened and deeply channeled distally; basal clusters on the surface of the adaxial lamina at the distal juxtacostal laminal cells rectangular, straight, with thin part of the leaf. The position of the species within the walls; basal marginal laminal cells rectangular, with thick genus is not clear. Although its leaf form and areolation transverse walls and thin lateral walls, hyaline in 3–4 demonstrate its similarity to G. tergestina (subg. rows; medial laminal cells short-rectangular, sinuose, Litoneuron), the longitudinal plicae in the most distal slightly thick-walled; distal laminal cells 1-stratose with leaves align it with G. caespiticia (subg. Guembelia). 2-stratose ridges, in apex entirely 2-stratose. Gemmae Sinuose laminal cell walls, characteristic of Grimmia, are clusters of globular, brown, multicellular gemmae only weakly present. abundantly present on the adaxial side of the distal lamina. Sexual condition probably dioicous, gametangia not seen. Capsule unknown.

3. COSCINODON Sprengel, Anleit. Kenntn. Gew. 3: 281. 1804 · [Greek koskinon, sieve, and odon, tooth, alluding to peristome]

Roxanne I. Hastings

Plants 4.5–15 mm, in dense cushions to loose mats, olivaceous to dark black-green. Leaves oval to ovate-lanceolate, keeled distally, margins plane, incurved, or recurved, distal lamina 1- or 2-stratose, specialized laminal and marginal chlorophyllose structures absent, usually long- awned; basal cells usually rectangular with straight walls; mid leaf and distal cells quadrate to short rectangular, straight or sinuose, usually thick-walled. Gemmae absent. Sexual condition autoicous or dioicous; perichaetial leaves enlarged. Seta short to long, straight. Capsule erect, immersed to exserted, symmetric, campanulate, ovoid to cylindrical, rarely cupulate; annulus poorly differentiated; operculum conic to rostrate, falling detached from the columella. Calyptra 1 campanulate, becoming cucullate with age, erose at base, large, covering /2 to all of capsule, plicate. Species 10 (5 in the flora): North America, South America, Eurasia. Species of Coscinodon are widespread across North America but are largely absent from the interior Great Plains and along the west coast. They all prefer dry acidic rocks. The largely calcareous regions of the continental interior do not provide suitable habitat for these species. Historically, North American authors have often considered Coscinodon to be a subgenus of Grimmia. Not until the 1980s did they follow the European approach distinguishing the two genera, the separation of which is based largely on the sporophyte. Coscinodon have large, campanulate, plicate calyptrae that commonly cover the capsule, whereas Grimmia have smaller, Coscinodon · GRIMMIACEAE 259 non-plicate calyptrae that are not campanulate. It is often stated that Coscinodon have very long awns compared to most Grimmia. This is true for the widespread species, C. calyptratus and C. cribrosus, and also C. yukonensis. But C. arctolimnius is muticous to short-awned and C. hartzii has awns not much longer than those of most Grimmia.

SELECTED REFERENCE Hastings, R. I. 1999. and biogeography of the genus Coscinodon (, Grimmiaceae) in North America, including a new species. Bryologist 102: 265–286.

1. Leaves non-plicate; lamina distally 1-stratose; leaf margins plane or recurved on one side at midleaf. 2. Autoicous; capsule exserted; peristome solid; leaves 1.4–2.4 mm, narrowly ovate- lanceolate, margins plane or recurved on one side at midleaf; western North America ...... 1. Coscinodon calyptratus 2. Dioicous; capsule immersed; peristome scalloped; leaves 0.9–1.2 mm, broadly ovate- lanceolate, margins plane; northern British Columbia to Alaska ...... 2. Coscinodon yukonensis 1. Leaves plicate at least in distal lamina; lamina distally 2-stratose; leaf margins incurved distally. 3. Capsule exserted; peristome present, xerocastique; leaf plications never to base; arctic ...... 5. Coscinodon hartzii 3. Capsule immersed; peristome present and hygrocastique or absent; leaf plications variable; temperate to arctic. 4. Peristome present; leaves never cucullate, plications not always reaching base; frequently long-awned; plants dark olivaceous; temperate to arctic ...... 3. Coscinodon cribrosus 4. Peristome absent; leaf apex often cucullate if muticous, inrolled if awned; plications extending to base; often muticous or with short awns; plants dark black-green; Arctic ...... 4. Coscinodon arctolimnius

1. Coscinodon calyptratus (Drummond) C. E. O. Jensen outcrops; moderate to high elevations (300–3000 m); in N. C. Kindberg, Eur. N. Amer. Bryin. 2: 241. 1897 Alta., B.C.; Ariz., Calif., Colo., Idaho, Minn., Mont., Nev., N.Mex., Oreg., S.Dak., Utah, Wash., Wis., Wyo. Grimmia calyptrata Drummond, Coscinodon calyptratus is common and widespread Musc. Amer., 60. 1828; Coscinodon in the dry interior mountain areas of western North hookeri Hampe; G. columbica America. To the east it is largely bounded by the front Kindberg ranges of the Rocky Mountains and acidic outliers such

5 5 as the Black Hills of South Dakota. Two disjunct sites in

5 Plants 7–10 mm, olivaceous. 5 5 Wisconsin and Minnesota define the eastward extent of 5 5 5 5 5 5 5 Leaves ovate to ovate-lanceolate, 5 5 the species. It is not found along the west coast 5 5 1.4–2.4 × 0.4–0.7 mm, margins mountains. Specimens from central Asia and New plane or one margin recurved at Zealand attributed to this North American endemic by mid leaf, apex plane, awn 0.4–1.4 J. Muñoz (1998c) instead represent an undescribed mm, lamina non-plicate; basal juxtacostal laminal cells species of Coscinodon and a Grimmia. The non-plicate, long-rectangular, 15–80 × 7–10 µm, evenly thin-walled; 1-stratose leaves with plane or one revolute margin, and basal marginal laminal cells quadrate to long-rectangular, exserted capsule clearly separate this species from all other 16–60 × 7–15 µm, thin or thick end walls and thin lateral North American Coscinodon. The autoicous sexual walls; medial laminal cells 1-stratose; distal laminal cells condition is unique in the genus but often difficult to 1-stratose. Sexual condition autoicous. Seta 1.4–2.4 mm. ascertain on any particular specimen. This species is more Capsule exserted, cylindric, commonly constricted at rim; commonly confused with G. longirostris (formerly G. peristome present, solid, xerocastique. affinis) than with any other Coscinodon. Both species Most common on dry sandstone and granitic boulders are autoicous, have a similar robust habit, have leaves and bedrock exposures but also limestone and volcanic that are strongly keeled and may have recurved margins, 260 GRIMMIACEAE · Coscinodon

COSCINODON

and have long-exserted capsules. If fertile and with walled; medial laminal cells 1-stratose; distal laminal cells capsules, the species are easily distinguished, as C. 1-stratose. Sexual condition dioicous. Seta 0.3–0.5 mm. calyptratus has large, plicate calyptra and has a thin- Capsule immersed, cupulate; peristome present, walled filmy annulus. In contrast, G. longirostris has scalloped, xerocastique. smaller, smooth calyptra and a large, prominent annulus. Siliceous or granite-slate outcrops; moderate to high Gametophytically, C. calyptratus has 1-stratose elevations (1500–1700 m); B.C., Yukon; Alaska. areolation across the distal and medial lamina except for Coscinodon yukonensis is known only from a 2-stratose margin. Its basal cells tend to be evenly thin- northwestern North America. The specimens were all walled. Grimmia longirostris has a 2-stratose distal originally identified as C. cribrosus, which it resembles lamina and a wide 2-stratose margin at mid leaf. Its basal in its immersed capsules and superficially cribrose cells tend to be thicker walled. peristome teeth. However, C. yukonensis is separated by its xerocastique peristome (i.e., opens when dry) and 2. Coscinodon yukonensis Hastings, Bryologist 102: non-plicate, plane margined, 1-stratose leaves. 278, figs., 42–51, 53–54. 1999 Additionally, unlike those of C. cribrosus, the peristome Plants 7–10 mm, dark brown teeth are not perforated; rather, the ornamentation has green. Leaves broadly ovate- been “scalloped out” down to the cell walls on either 5 lanceolate, 0.9-1.2 × 0.4 mm, side of the teeth. Thus, while superficially appearing 5 margins plane, apex plane, awn perforated, the cell walls are intact. Its leaves resemble 5 0.2–1 mm, lamina non-plicate; those of C. calyptratus but are much smaller, more basal juxtacostal laminal cells broadly ovate, and never revolute. Additionally, it has rectangular to long-rectangular, an immersed capsule and the peristome teeth are 27–51 × 8–12 µm, thin- to thick- scalloped, not solid as in C. calyptratus. The basal walled; basal marginal laminal marginal laminal cells are oblate to quadrate, unlike those cells oblate to quadrate, 5–12 × 9–12 µm, evenly thick- of any other Coscinodon in North America. Coscinodon · GRIMMIACEAE 261

3. Coscinodon cribrosus (Hedwig) Spruce, Ann. Mag. 4. Coscinodon arctolimnius (Steere) Steere, J. Hattori Nat. Hist., ser. 2, 3: 491. 1849 Bot. Lab. 42: 446. 1977 (as arctolimnia) Grimmia cribrosa Hedwig, Sp. Grimmia arctolimnia Steere,

5 Musc. Frond., 76. 1801; Bryologist 77: 230, figs. 1–19. 1974

5 Coscinodon persoonii Hampe; 5 Plants 5–10 mm, dark black- 5 5 C. pulvinatus Sprengel 5 green. Leaves ovate-lanceolate, 5 5 5 5 5 5 Plants 4.5–7 mm, dark olivaceous. (0.8–)1.5–1.8 × (0.3–)0.6–0.8 mm, 5 5 5 Leaves oval to ovate-lanceolate, margins incurved distally, apex 5

5 5 5

5 1.1–1.9 × 0.4–0.7 mm, margins often cucullate, awn 0.3–0.5 mm incurved distally, apex plane, awn or absent, lamina 2-plicate, 0.1–1.1 mm, lamina 2-plicate, extending from apex to base; basal plications not always reaching base; basal juxtacostal juxtacostal laminal cells short-rectangular, 19–33 × 9– laminal cells quadrate to long-rectangular, 20–47 × 8– 13 µm, evenly thick-walled; basal marginal laminal cells 12 µm, evenly thick-walled; basal marginal laminal cells quadrate to short-rectangular, 13–27 × 9–13 µm, evenly quadrate to rectangular, 12–34 × 6–12 µm, thin or thick thin-walled; medial laminal cells patchy 2-stratose; distal end walls and thin lateral walls; medial laminal cells 1- laminal cells 2-stratose. Sexual condition dioicous. Seta stratose; distal laminal cells 2-stratose. Sexual condition 0.3–0.5 mm. Capsule immersed, ovoid to campanulate; dioicous. Seta 0.8–1.2 mm. Capsule emergent, ovoid to peristome absent. campanulate; peristome present, cribrose, hygrocastique. Mineralized siliceous rock; low elevations (160 m); Acidic, dry sandstone, shale, and granitic boulders and N.W.T.; Asia (Mongolia, Pakistan). bedrock exposures but also found on volcanic outcrops Until 1996, Coscinodon arctolimnius was considered and granodiorites; low to high elevations (0–3300 m); to be endemic to the eastern shore of Great Bear Lake in Greenland; Alta., B.C., Nfdl. and Labr., N.W.T., Nunavut, the Northwest Territories of Canada. When he first Ont., Que., Yukon; Alaska, Ariz., Colo., Maine, N.Mex., described it, Steere suggested that the species evolved in N.Y., S.Dak., Tenn., Tex.; Eurasia. that site as a result of radiation-induced genetic mutations Coscinodon cribrosus is an uncommon known in a population of C. cribrosus. However, with the from widely disjunct sites across North America. It has discovery of a new subspecies (R. I. Hastings and H. been suggested that this distribution is due to a restricted Deguchi 1997), from both Pakistan and Mongolia, it occurrence on mineral-rich rocks. The sites from appears that the species has a much wider distribution. Ellesmere Island, Tennessee, Colorado, South Dakota, Although undoubtedly closely related to C. cribrosus, Thunder Bay, and Alaska support this contention, as they there is now no reason to assume that C. arctolimnius are all reported to be either heavy-metal- or copper- evolved in North America. It is separated from other bearing deposits or the specimens are associated with the North American Coscinodon by its green-black color, “copper-moss,” Mielichhoferia mielichhoferi. However, cucullate leaf apex (incurved if awned), and absence of a specimens from other areas have been collected on peristome. sandstones and shales. The distribution appears to have SELECTED REFERENCE Hastings, R. I. and H. Deguchi. 1997. been largely controlled by the extent of the Cretaceous Taxonomy of Coscinodon arctolimnius (Bryopsida, Grimmiaceae) epicontinental seaway that flooded much of central including a new subspecies from Asia. Bryologist 100: 50–55. North America from the Gulf of Mexico to the Yukon (R. I. Hastings 1999). The species does not occur within 5. Coscinodon hartzii C. E. O. Jensen, Meddel. the boundaries of the seaway except on isolated Grønland 15: 422, figs. 1–8. 1897 granodiorite outcrops that were exposed subsequent to Plants 10–15 mm, pale olivaceous. the retreat of the seaway. Coscinodon cribrosus is 5 Leaves oval to ovate-lanceolate, recognized by its 2-plicate, 2-stratose leaves with incurved 5 1.1–1.6 × 0.3–0.6 mm, margins 5 margins and emergent capsule with a well-developed incurved distally, apex plane, awn cribrose peristome. It is the only species of the genus 0.1–0.6 mm, lamina 2-plicate, that has a hygrocastique peristome, i.e., its peristome plications never extended to base; opens when wet and is closed when dry. basal juxtacostal laminal cells short- to long-rectangular, 21–55 × 9–12 µm, evenly thick-walled; basal marginal laminal cells quadrate to long-rectangular, 12–51 × 9–15 µm, thin or thick end walls and thin lateral walls; medial laminal cells 1-stratose; distal laminal cells 2–3-stratose. Sexual condition dioicous. Seta 1.5–1.8 mm. Capsule exserted, cylindrical; peristome present, cribrose, xerocastique. 262 GRIMMIACEAE · Coscinodon· Jaffueliobryum

COSCINODON ° JAFFUELIOBRYUM

Shale, slate, and granitic rock in tundra; moderate to W. C. Steere’s (1978) and J. Muñoz’s (1998c) citations of high elevations (1000–1700 m); Greenland; Yukon; C. calyptratus from those sites have turned out to be Alaska. C. hartzii. A new location in southeastern Yukon suggests Coscinodon hartzii is endemic to arctic regions of that this species may be widespread across northern North America. Until 1992, it was known only from its North America. Coscinodon hartzii is recognized by its type locality at Scoresby Sound in eastern Greenland. weakly 2-plicate leaves coupled with an exserted capsule B. M. Murray (1992) reported collections from Mt. and cribrose peristome. It commonly also has 3-stratose Michelson and the Philip Smith Mountains of Alaska. regions in the distal parts of its stem leaves.

4. JAFFUELIOBRYUM Thériot, Rev. Bryol., n. s. 1: 192, plate 8, figs. 1–5. 1928 · [For Félix Jafuell, 1857–1931, clergyman who collected plants in South America, and Greek bryum, moss]

John R. Spence

Plants 3–15(–20) mm, in dense cushions, dark olive, yellow-green, or dull green to olivaceous. Leaves broadly ovate to obovate, concave or somewhat keeled, margins plane, smooth or weakly serrate to erose distally, distal lamina mostly 1-stratose, rarely with 2-stratose streaks, specialized laminal and marginal chlorophyllose structures absent, awn short to twice lamina length, distal leaves typically with longer awns; basal cells elongate-rectangular to oblong, with straight, somewhat lax, sometimes weakly hyaline walls; mid leaf and distal cells isodiametric, oval to irregularly rhomboidal; distal cells becoming hyaline with age. Gemmae absent. Sexual condition Jaffueliobryum · GRIMMIACEAE 263 autoicous or rarely cryptoicous; perichaetial leaves enlarged, long-awned. Seta short, straight. Capsule erect, immersed, symmetric, globose to ovoid; annulus differentiated, typically persistent; operculum conic, short to long-rostrate, falling detached from columella. Calyptra campanulate, mitrate, sometimes erose or lobed at base, large, covering 1/2–3/4 of capsule, plicate. Species 3 (2 in the flora): North America, Mexico, South America (Bolivia), c Eurasia (China, Mongolia, Russia). Jaffueliobryum is distributed in cold to hot and arid regions, and usually is found on calcareous rock. It appears to be closest to Coscinodon, from which it differs by the broader, less strongly keeled, mostly 1-stratose leaves, strictly autoicous-cryptoicous sexuality, and calciphilous ecology. The third species, J. arsenei (Thériot) Thériot, larger than our species, is endemic to Mexico and is characterized by a long-rostrate operculum and an exserted capsule.

SELECTED REFERENCE Churchill, S. P. 1987. Systematics and biogeography of Jaffueliobryum (Grimmiaceae). Mem. New York Bot. Gard. 45: 691–708.

1. Distal lamina acute to acuminate, leaves distinctly keeled, proximal stem leaves mostly spreading ...... 1. Jaffueliobryum raui 1. Distal lamina broadly acute-rounded, leaves not keeled, proximal stem leaves mostly appressed...... 2. Jaffueliobryum wrightii

1. Jaffueliobryum raui (Austin) Thériot, Rev. Bryol., n. s. sandstone or limestone rock, open arid to semi-arid shrub, 1: 193. 1928 woodland communities, grasslands, rarely on compacted Grimmia raui Austin, Bull. Torrey sandy soil; moderate to high elevations (200–2100 m); Bot. Club 6: 46. 1875 Alta.; Ariz., Calif., Colo., Iowa, Kans., Minn., Mont., Nebr., Nev., N.Mex., Okla., S.Dak., Tex., Utah, Wis. Plants in small dense cushions or Jaffueliobryum raui is a common species in drier parts turfs, yellow-green to dark of the United States, especially on the Great Plains and 5 olivaceous, hoary. Stems 5–20

5 the Colorado Plateau, extending to the Mohave Desert 5 5 5

5 5 5 mm, sparsely branched. Leaves 5 5 5 5 of California. This species appears to be disjunct to 5 5 5 crowded, ovate to obovate, 5 southern Alberta, but the distribution is likely continuous, imbricate to appressed-julaceous as the band of calcareous bedrock on which it occurs distally, somewhat spreading to runs along the Rocky Mountain Front Range from squarrose-recurved proximally, 0.6–1.2 mm excluding Alberta well into Montana. Sites in the Driftless Area of awn, apex acute to acuminate, lamina 1-stratose to rarely Iowa, Minnesota, and Wisconsin may be truly disjunct. 2-stratose in bands, awn length highly variable, 0.3–1.4 However, more collecting is likely to fill in gaps in its mm, hyaline; costa in transverse-section distinctly keeled; range in the northern Great Plains and clarify its status proximal cells rectangular, 15–40 × 10–20 µm, often in the Great Basin. appearing lax; mid leaf cells isodiametric to short-oval, Jaffueliobryum wrightii is very closely related, and the (5–)8–12(–20) µm; distal cells somewhat longer than mid two species have been treated as one for much of their leaf cells. Sexual condition autoicous; perichaetial leaf taxonomic history. In addition to traits used in the key, lamina to 1.5 mm, awn 1–1.7 mm. Seta 0.4–0.6 mm. there are a variety of other partially overlapping Capsule yellow-brown turning red-brown with age, ovoid characters that separate the two, including leaf width at to subglobose, 0.8–1 mm; operculum short-rostrate, 0.5– widest point, lamina length, costal width, lamina cell 0.6 mm. length and angle between the margin at the leaf widest Capsules mature spring–summer depending on point and the tip of the awn, and capsule and operculum elevation. Widespread and locally common on dry length.