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flat-rock grimmia, oval dry rock moss

grimmia dry rock moss

Habit Plants in loose tufts, dark green to brownish black. Plants 5–40(–70) mm, in dense cushions to loose mats, olivaceous, dark black-green to rusty-red-brown.
Stems

1–3 cm.

Gemmae

present or absent.

Leaves

ovate-lanceolate from an ovate base, 1.7–4 × 0.4–0.8 mm, both margins plane, incurved distally, intermarginal bands absent, awn 0.5–1 mm, not decurrent, typically narrowly attached, acute, costa narrow proximally;

basal juxtacostal laminal cells usually elongate (sometimes short-rectangular), usually sinuose, and usually with thick lateral walls;

basal marginal laminal cells quadrate to long-rectangular, straight, with thick transverse and thin lateral walls, green, not to distinctly hyaline;

medial laminal cells rounded to quadrate, straight, thick-walled;

distal laminal cells 2-stratose, quadrate, thick-walled.

broadly oblong-ovate, oblong-lanceolate, to narrowly ovate-lanceolate, rarely ligulate, concave or keeled distally, margins plane, incurved or recurved, distal lamina 1-stratose to multistratose, specialized laminal and marginal chlorophyllose structures absent, muticous to long-awned but awns only rarely longer than lamina;

basal cells oblate to elongate, with straight or sinuose and thin to thick cell walls;

mid leaf and distal cells quadrate to rectangular, usually sinuose and thick-walled.

Seta

straight, 4–6 mm.

short to long, straight, arcuate, or rarely sigmoid.

Sexual condition

autoicous or dioicous;

perichaetial leaves enlarged or not.

Capsule

occasionally present, exserted, yellow-brown, oblong-ovoid, exothecial cells short-rectangular, thin-walled, stomata present, annulus of 2–3 rows of rectangular, thick-walled cells, operculum long-rostrate, peristome solid, split in distal half.

erect, rarely pendent, immersed to long exserted, symmetric or rarely ventricose (subg. Grimmia), ovoid to obloid, rarely globose or cylindric;

annulus poorly differentiated or well-defined comprised of quadrate, thin-walled or quadrate to rectangular, thick-walled cells;

operculum mammillate, conic, or rostrate, falling detached from the columella.

Calyptra

cucullate.

mitrate or cucullate, not erose, small to medium, usually covering 1/2 or less of capsule, sometimes just covering operculum, smooth.

Perichaetial

leaves enlarged.

Grimmia ovalis

Grimmia

Habitat Dry, exposed to partially shaded, acidic, sandstone, granite and basalt, montane to alpine
Elevation (low to) moderate to high elevations (30-)1000-2500 m ((low to) moderate to high elevations (100-)3300-8200 ft)
Distribution
from FNA
AZ; CA; CO; ID; MN; MO; MT; NM; NV; NY; OR; PA; TX; UT; VA; WA; WY; AB; BC; ON; QC; YT; Greenland; Eurasia; Africa (Algeria, Morocco)
[WildflowerSearch map]
from USDA
North America; Mexico; Central America; South America; Eurasia; Africa; Pacific Islands; Australia; Antarctic
Discussion

Grimmia ovalis is common and widespread in high elevation sites in western North America from southern British Columbia along the Rocky Mountain corridor to southern New Mexico and south central California. H. A. Crum and L. E. Anderson (1981) rejected all reports of G. ovalis from eastern North America. However, we have seen specimens from that area, although scattered and rare. It is not surprising that the species occurs in the eastern part of the continent given that it is widespread across the Laurasian continental plates and in India. Outlier sites in the Yukon and southern Greenland connect the North American populations to those in Asia and Europe, respectively. In western North America, G. ovalis is most often confused with G. longirostris and G. laevigata. While superficially similar to G. longirostris, G. ovalis has concave leaves with plane margins and is dioicous, while G. longirostris has keeled leaves with a recurved margin and is autoicous. Other points of separation are discussed under G. longirostris. Typical specimens of G. ovalis are readily separated from G. laevigata by their ovate-lanceolate leaves with a well-defined ovate base and narrowly attached awns. However, both of these species are variable in leaf shape, and while the awn of G. ovalis is usually narrowly attached, sometimes it is quite broad and may border on being decurrent. These specimens can be identified by the width of the costa and the basal areolation. Grimmia ovalis has a costa that is narrow at the base, while G. laevigata has a distinctly broad costa covering up to 1/3 of the base. The basal marginal cells of G. ovalis are most often short- to long-rectangular and hyaline while those of G. laevigata are always oblate to quadrate and not hyaline. In eastern North America, specimens of G. ovalis have been misidentified as G. olneyi. Aside from seta and capsule differences, G. ovalis has rectangular basal marginal laminal cells and its basal juxtacostal cells are long-rectangular to elongate; see also under G. olneyi.

Based on identifications by Greven, W. A. Weber et al. (2003) reported Grimmia bernoullii in the United States. That species differs from G. ovalis by its more ovate leaves without shoulders and with plane margins, costa broad at the base and disappearing in mid leaf, and sporophytically by its ellipsoid capsule with long-rostrate, straight operculum, and mitrate calyptra. Hastings has examined duplicate specimens from Missouri of those cited by Weber et al. and has determined that they do not deviate significantly from G. ovalis, having rather narrow leaves, costa narrow at the base and remaining strong in mid leaf, and with many leaves having incurved margins. The specimens were sterile and therefore sporophytic characters could not be determined. Based on these observations Hastings excludes G. bernoullii from the North American flora, although Greven would still retain the species.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Species 95 (43 in the flora).

The genus Grimmia is found on all continents. However, nearly half (44) of the species are endemics and have restricted distributions. Most species of Grimmia prefer dry and temperate or cold environments—there is no species only known from tropical areas. Nearly all species of Grimmia are saxicolous with a marked preference for acidic bedrock. Only the subgenus Grimmia prefers calcareous rock, with a few other species in other subgenera characteristic of neutral to basic substrates.

Hastings has attempted to place species in the subgenera Grimmia, Guembelia, and Litoneuron into groups of related or similar-looking ones. Because of the diversity within subg. Rhabdogrimmia, Greven felt it most convenient to present this group largely in alphabetical order. While the proper subdivision of Grimmia remains uncertain, both authors agree that for purposes of identification, the present division is best for this very complicated genus. Hastings authored the accounts for species 1–8, 10, 11, 13–16, 18, and 20–24, Greven those for species 10, 13, 18, 20, 25–43. Both authors together wrote the entire key.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Costa ending well before the apex; leaf tip rounded to cucullate, muticous; lamina uniformly 1-stratose; laminal cells all quadrate.
G. mollis
1. Costa reaching apex; leaf tip rounded to acute, muticous or awned; lamina usually with 2-stratose areas towards margins or with 2-stratose ridges; laminal cells oblate, quadrate, rectangular or elongate
→ 2
2. Seta eccentrically attached to capsule base; capsule immersed, smooth, ventricose; stomata 3-4, large, at base of capsule (subg. Grimmia)
→ 3
2. Seta centrally attached to capsule base; capsule immersed to exserted, smooth to plicate, not ventricose; stomata none to many, small, at neck to base of capsule
→ 6
3. Distal leaves concave-keeled; distal lamina 1-stratose or with 2-stratose patches, margins 1- or 2-stratose; annulus absent or reduced to 1-2 rows of small cells; operculum mammillate
→ 4
3. Distal leaves concave; distal lamina 2-stratose with 2-stratose margins; annulus prominent; operculum rostrate
→ 5
4. Peristome present, annulus absent; distal lamina 1-stratose, margins 1- or 2- stratose
G. plagiopodia
4. Peristome absent, annulus present; distal lamina 1-stratose with 2-stratose patches, margins 2-stratose.
G. anodon
5. Basal laminal cells thick-walled; gonioautoicous; peristome fully developed.
G. americana
5. Basal laminal cells thin-walled; dioicous; peristome rudimentary.
G. crinitoleucophaea
6. Leaves merely 1-stratose distally; margins at most narrowly 2-stratose; one or both leaf margins recurved (G. reflexidens plane) (mostly subg. Rhabdogrimmia)
→ 7
6. Leaves 2-3-stratose distally or 1-stratose with distal margins widely 2-stratose (G. sessitana and some G. donniana); leaf margins incurved, plane or recurved
→ 26
7. Costa projecting on abaxial side with two wings.
G. ramondii
7. Costa on abaxial side smooth or angled
→ 8
8. Leaves muticous, without awns or only distal leaves with very short awn or hyaline tip
→ 9
8. At least distal or perichaetial leaves with distinct awns
→ 13
9. Leaves cucullate.
G. atrata
9. Leaves not cucullate
→ 10
10. Leaves squarrose when moist; medial laminal cells oblate to rounded quadrate
G. lisae
10. Leaves erect to patent when moist; medial laminal cells quadrate to short-rectangular
→ 11
11. Basal marginal cells with thickened transverse walls; seta arcuate; globular gemmae occasionally present on leaves.
G. trichophylla
11. Basal marginal cells with thin transverse walls; seta straight or slightly arcuate; gemmae absent
→ 12
12. One leaf margin recurved; medial laminal cells sinuose; seta straight or slightly arcuate
G. elongata
12. Leaf margins both recurved; medial laminal cells with nodulose walls; seta straight.
G. lesherae
13. Awns nearly equal to or much longer than lamina, decurrent, seta straight to flexuose
→ 14
13. Awns typically shorter than lamina (G. pulvinata can be long); decurrent or not; seta arcuate to cygneous
→ 15
14. Awns 1-2 mm, leaf margins plane, [seta straight]; only known from Maritime provinces in North America.
G. reflexidens
14. Awns 2-4 mm, leaf margins both recurved; seta flexuose; endemic to Montana and Idaho.
G. brittoniae
15. Leaves abruptly narrowed to awns, apex rounded; autoicous
→ 16
15. Leaves gradually tapering to awns, apex narrowed; dioicous
→ 18
16. Operculum rostrate; basal marginal laminal cells quadrate to short-rectangular
G. pulvinata
16. Operculum mammillate to conical; basal marginal laminal cells short- to long-rectangular
→ 17
17. Awns only on perichaetial leaves; basal juxtacostal cells with thin, straight walls; marginal distal lamina 2-stratose; acidic rocks.
G. moxleyi
17. Awns on all distal leaves; basal juxtacostal cells with thick, nodulose walls; marginal distal lamina 1-stratose; basic rocks.
G. orbicularis
18. Clusters of gemmae easily visible on abaxial side of leaf lamina or on leaf tips
→ 19
18. Clusters of gemmae absent or in obscure clusters in leaf axils
→ 21
19. Multicellalar gemmae on abaxial side of leaf lamina in distal leaves; leaves crisped and contored.
G. torquata
19. Cluster of gemmae only on leaf tips; leaves straight
→ 20
20. Leaves with blunt eroded apices; distal cells with pseudo- papillose walls; stem central strand present.
G. anomala
20. Leaves with acuminate apices; distal cells with smooth walls; stem central strand absent.
G. hartmanii
21. Leaves narrowly ovate-lanceolate to linear-lanceolate, distinctly incurved or contorted when dry.
G. incurva
21. Leaves lanceolate to ovate-lanceolate, straight, appressed, only occasionally somewhat contorted when dry
→ 22
22. Central strand absent; basal juxtacostal leaf cells uniformly elongate to linear, strongly nodulose and thick-walled; mid leaf cells extremely sinuose and very thick-walled; leaf margins both recurved
→ 23
22. Central strand present; basal juxtacostal cells short- to long-rectangular, elongate cells absent or few and scattered; mid leaf cells straight to sinuose, thin- to thick-walled; leaf margins one or both recurved
→ 24
23. Costa circular, projecting in transverse section; basal marginal cells thin-walled; distal leaves straight, awns not decurrent.
G. attenuata
23. Costa semicircular, not projecting in transverse section; basal marginal cells thick-walled; distal leaves often secund, awns often decurrent
G. leibergii
24. Leaves squarrose when moist; mid leaf cells rounded to oblate, straight.
G. lisae
24. Leaves erect to patent when moist; mid leaf cells rounded-quadrate to short-rectangular, sinuose
→ 25
25. Blackish green tufts; costa angled or bluntly winged in transverse section; awns denticulate, occasionally decurrent; capsule globose, brown.
G. muehlenbeckii
25. Yellowish green tufts; costa semicircular in transverse section; awns smooth to slightly denticulate, not decurrent; capsule oblong-ovoid, yellow-green
G. trichophylla
26. Leaves concave; costa not prominent; margins plane or incurved; dioicous (subg. Litoneuron)
→ 27
26. Leaves keeled; costa prominent; margins recurved, plane or incurved; autiocous or dioicous (mostly subg. Guembelia)
→ 33
27. All leaves muticous, straight or falcate
→ 28
27. At least distal leaves awned, straight
→ 29
28. Leaves oblong-lanceolate to ligulate, straight, cucullate, obtuse-rounded; basal marginal laminal cells short-rectangular; widely distributed.
G. unicolor
28. Leaves oblong-lanceolate, homomallous-falcate, subulate, uncinate; basal marginal laminal cells quadrate to short-rectangular; endemic to Pacific Coast region.
G. hamulosa
29. Leaves oblong-ovate to oblong-lanceolate; basal marginal laminal cells oblate to quadrate; costa broad at base; awn broadly attached and decurrent.
G. laevigata
29. Leaves ovate-lanceolate from an ovate base; basal marginal laminal cells quadrate to long-rectangular; costa narrow at base; awn typically narrowly attached and not decurrent
→ 30
30. Costa-like multistratose leaf bands present in transverse section.
G. serrana
30. Costa-like multistratose leaf bands absent
→ 31
31. Capsule emergent, gymnostomous, stomata absent, operculum mammilate; basal laminal cells uniformly short-rectangular, straight and thin-walled.
G. nevadensis
31. Capsule exserted, peristome fully developed, stomata present, operculum rostrate; basal marginal and juxtacostal laminal cells typically contrasting in length or thickness
→ 32
32. Basal marginal laminal cells quadrate; basal juxtacostal laminal cells quadrate to short-rectangular; seta sigmoid; capsule wrinkled when dry; perichaetial leaves not enlarged; endemic to eastern North America.
G. olneyi
32. Basal marginal laminal cells quadrate to long-rectangular; basal juxtacostal laminal cells rectangular to elongate; seta straight; capsule smooth when dry; perichaetial leaves enlarged; widely distributed.
G. ovalis
33. Margins plane or incurved
→ 34
33. Margins recurved on one or both sides
→ 44
34. Costa transverse section circular distally; awn hyaline-tipped to short (0.3 mm), decurrent
G. teretinervis
34. Costa transverse section semicircular distally; awn short to long, usually prominent, decurrent or not
→ 35
35. Seta arcuate; leaves spirally curved around stem when dry, flagelliform innovations present
G. funalis
35. Seta straight; leaves straight when dry, flagelliform innovations absent
→ 36
36. Gemmae abundant on adaxial distal leaf surface.
G. shastae
36. Gemmae absent on leaves
→ 37
37. Distal laminal cells bulging
→ 38
37. Distal laminal cells not bulging
→ 40
38. Leaf margins plane; cladautoicous; basal juxtacostal laminal cells rectangular to elongate; distal juxtacostal laminal cells 1-stratose; stomata present.
G. sessitana
38. Leaf margins incurved; dioicous; basal juxtacostal laminal cells quadrate to short-rectangular; distal juxtacostal laminal cells 2-stratose; stomata present or absent
→ 39
39. Stomata absent; leaves not plicate, not cucullate.
G. alpestris
39. Stomata present; leaves weakly to rarely strongly plicate distally, cucullate
G. caespiticia
40. Margins plane throughout; basal marginal laminal cells short- to long-rectangular; stomata present; autoicous
→ 41
40. Margins incurved distally; basal marginal laminal cells quadrate to short-rectangular; stomata absent; dioicous
→ 42
41. Distal juxtaxcostal laminal cells 2-stratose, occasionally 1-stratose; basal marginal leaf cells long-rectangular, hyaline; annulus large, of 2 rows of rectangular cells, revoluble; calyptra mitrate.
G. donniana
41. Distal juxtacostal laminal cells 1-stratose; basal marginal leaf cells short- to long-rectangular, rarely hyaline; annulus small, of 1 row of quadrate cells, not revoluble; calyptra cucullate.
G. sessitana
42. Capsule immersed to emergent, wide-mouthed, peristome rudimentary; endemic to California.
G. mariniana
42. Capsule exserted, narrow-mouthed, peristome present, fully developed; widespread in western North America
→ 43
43. Basal juxtacostal laminal cells short- to long-rectangular, distinct from quadrate to short-rectangular basal marginal cells; medial laminal cells rounded, thick-walled.
G. montana
43. Basal juxtacostal and marginal cells quadrate to short-rectangular, basal areolation relatively uniform; medial laminal cells quadrate to short-rectangular, thin-walled.
G. alpestris
44. Leaves muticous, cucullate.
G. atrata
44. Leaves awned, not cucullate
→ 45
45. Leaves spirally curved around stem when dry, flagelliform innovations present.
G. funalis
45. Leaves straight when dry, flagelliform innovations absent
→ 46
46. Sporophytes present
→ 47
46. Sporophytes absent
→ 51
47. Capsule immersed
→ 48
47. Capsule exserted
→ 49
48. Stem central strand present, epidermis thin; leaf margins 2-stratose, not thickened, one margin recurved (rarely both); leaves broadly ovate- lanceolate.
G. arizonae
48. Stem central strand absent, epidermis thick; leaf margins 3(-4) stratose, thicker than juxtacostal lamina, usually both margins recurved; leaves narrowly lanceolate from an ovate base.
G. pilifera
49. Seta arcuate; dioicous.
G. elatior
49. Seta straight; autoicous
→ 50
50. Annulus small, 1 row of quadrate cells; stomata in 1 row; basal juxtacostal cells straight and thin-walled.
G. sessitana
50. Annulus prominent, 2 rows of rectangular cells; stomata in 2-3 rows; basal juxtacostal cells sinuose and thick walled.
G. longirostris
51. Basal juxtacostal cells straight, thin-walled; distal juxtacostal lamina 1-stratose, cells often bulging; plants small (less than 1 cm), blackish; moist, alpine habitats.
G. sessitana
51. Basal juxtacostal cells sinuose, thick-walled; distal juxtacostal lamina at least 2-stratose, cells not bulging; plants robust (greater than 1cm), yellow-green to very dark olivaceous; dry, widely distributed
→ 52
52. Leaf margins 2-stratose, not thickened; stem central strand present, epidermis thin
→ 53
52. Leaf margins multistratose and thickened; stem central strand absent, epidermis thick
→ 54
53. Autoicous; costa transverse section reniform; leaves sheathing; widely distributed.
G. longirostris
53. Dioicous; costa transverse section semicircular; leaves not sheathing; American Southwest
G. arizonae
54. Leaves narrowly lanceolate from an ovate base, usually narrowly recurved on both margins; distal lamina without multistratose bands, never papillose.
G. pilifera
54. Leaves broadly lanceolate, broadly recurved on one margin; distal lamina with multistratose bands, occasionally papillose
G. elatior
Source FNA vol. 27, p. 244. FNA vol. 27, p. 225. Authors: Roxanne I. Hastings, Henk C. Greven.
Parent taxa Grimmiaceae > subfam. Grimmioideae > Grimmia > subg. Litoneuron Grimmiaceae > subfam. Grimmioideae
Sibling taxa
G. alpestris, G. americana, G. anodon, G. anomala, G. arizonae, G. atrata, G. attenuata, G. brittoniae, G. caespiticia, G. crinitoleucophaea, G. donniana, G. elatior, G. elongata, G. funalis, G. hamulosa, G. hartmanii, G. incurva, G. laevigata, G. leibergii, G. lesherae, G. lisae, G. longirostris, G. mariniana, G. mollis, G. montana, G. moxleyi, G. muehlenbeckii, G. nevadensis, G. olneyi, G. orbicularis, G. pilifera, G. plagiopodia, G. pulvinata, G. ramondii, G. reflexidens, G. serrana, G. sessitana, G. shastae, G. teretinervis, G. torquata, G. trichophylla, G. unicolor
Subordinate taxa
G. alpestris, G. americana, G. anodon, G. anomala, G. arizonae, G. atrata, G. attenuata, G. brittoniae, G. caespiticia, G. crinitoleucophaea, G. donniana, G. elatior, G. elongata, G. funalis, G. hamulosa, G. hartmanii, G. incurva, G. laevigata, G. leibergii, G. lesherae, G. lisae, G. longirostris, G. mariniana, G. mollis, G. montana, G. moxleyi, G. muehlenbeckii, G. nevadensis, G. olneyi, G. orbicularis, G. ovalis, G. pilifera, G. plagiopodia, G. pulvinata, G. ramondii, G. reflexidens, G. serrana, G. sessitana, G. shastae, G. teretinervis, G. torquata, G. trichophylla, G. unicolor
Synonyms Dicranum ovale, G. commutata, G. ovata, G. ovata var. gracilis Dryptodon, Hydrogrimmia
Name authority (Hedwig) Lindberg: Acta Soc. Sci. Fenn. 10: 75. (1871) Hedwig: Sp. Musc. Frond., 75. 1801 ,
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