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BFNA Title: Grimmia |
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Grimmia - Grimmiaceae Edit Level Version 3 - 05513 plus revision by Hastings 9/05 and July 2006 Grimmia -
Grimmiaceae XXX.
GRIMMIA Hedwig, Spec. Musc.
Frond.: 75. 1801 * [For Dr. J.F.K. Grimm,
physician and botanist of Roxanne I. Hastings Henk C. Greven Dryptodon Bridel; Hydrogrimmia ( Plants 5--40(--70) mm, in tight cushions or
sometimes loose mats, olive green, dark black-green to rusty-red-brown. Stem leaves variable, 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 lacking, muticous to long-awned but awns only
rarely longer than the lamina, mid-leaf cells quadrate to rectangular,
usually with sinuose, thick walls, basal cells oblate to elongate, with
straight to sinuose and thin to thick cell walls. Gemmae present or absent. Sexual
condition autoicous or dioicous. Perichaetial
leaves enlarged or not. Seta
short to long, straight, curved, or rarely sigmoid. Capsule erect, rarely pendent, immersed to long exserted, symmetric or rarely ventricose (Gasterogrimmia), ovate to obloid,
rarely globose or cylindric, with a poorly differentiated annulus, or
well-defined annulus of quadrate, thin-walled or rectangular, thick-walled
cells, operculum mammillate, conic or rostrate, falling detached from the
columella. Calyptra mitrate or
cucullate, not erose, small to medium, usually covering 1/2 or less of
capsule, sometimes just covering operculum, smooth. Species 93 (43 in the flora): North
America, 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 Gasterogrimmia prefers calcareous
rock, with a few other species in other subgenera characteristic of neutral
to basic substrates. R.
I. Hastings has attempted to place species in the subgenera Gasterogrimmia, Guembelia and Litoneuron into groups of related or
similar-looking species. Because of the diversity within subg. Rhabdogrimmia, H. C. 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 was best for this very complicated genus. Note: R.I.H.
authored species accounts for 1--8, 10, 11, 13--16, 18, and 20--24; H.C.G.
authored species accounts 10, 13, 18, 20, 25--43. Both authors worked
together to write the entire key. SELECTED REFERENCES Deguchi, H. 1987. Studies on some
Peruvian species of Grimmiaceae. In; H. Inoue (ed.), Studies on Cryptogams in
1. Costa
ending well below the apex; leaf tip rounded to cucullate, muticous; lamina
uniformly 1-stratose; laminal cells all quadrate .…………………..… 42. Grimmia 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. Seta eccentrically attached to capsule base;
capsule immersed, smooth, ventricose; stomates 3--4, large, at base of
capsule ...……(subg. Gasterogrimmia) 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.
Peristome present, annulus absent; distal lamina 1-stratose, margins 1- or
2-stratose .............................1. Grimmia plagiopodia 4.
Peristome absent, annulus present; distal lamina 1-stratose with 2-stratose
patches, margins 2-stratose .............. 2. Grimmia anodon 3.
Distal leaves concave; distal lamina 2-stratose with 2-stratose margins;
annulus prominent; operculum rostrate. 5.
Basal laminal cells thick-walled; gonioautoicous; peristome fully developed
............................................ 3. Grimmia 5.
Basal laminal cells thin-walled; dioicous; peristome rudimentary
.................................................................. 4. Grimmia poecilostoma 2. Seta centrally attached to capsule base; capsule
immersed to exserted, smooth to plicate, not ventricose; stomates none to
many, small, at neck to base of capsule. 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. 7.
Leaves concave; costa not prominent; margins plane or incurved; dioicous
……………………………. (subg. Litoneuron) 8. All leaves muticous, straight or falcate. 9.
Leaves oblong-lanceolate to ligulate, straight, cucullate, obtuse-rounded;
basal marginal laminal cells short-rectangular; widely distributed
.……………………..…….. 18. Grimmia unicolor 9.
Leaves oblong-lanceolate, homomallous-falcate, subulate, uncinate; basal
marginal laminal cells quadrate to short-rectangular; endemic to 8. At least distal leaves awned, straight. 10.
Leaves oblong-ovate to oblong-lanceolate; basal marginal laminal cells oblate
to quadrate; costa broad at base; awn broadly attached and decurrent ..............................................
24. Grimmia laevigata 10.
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. 11.
Costa-like multistratose leaf bands present in transverse section……..…...…… 23. Grimmia
serrana 11.
Costa-like multistratose leaf bands absent. 12.
Capsule exserted, peristome fully developed, stomates present, operculum
rostrate; basal marginal and juxtacostal laminal cells typically contrasting
in length or thickness. 13.
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 ….…….
21. Grimmia ovalis 13. 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 ...............
20. Grimmia olneyi 12.
Capsule emergent, gymnostomous, stomates absent, operculum mammilate; basal
laminal cells uniformly short-rectangular, straight and thin-walled ....................
22. Grimmia nevadensis [7 shifted to left ] 7.
Leaves keeled; costa prominent; margins recurved, plane or incurved;
autiocous or dioicous .....................………….… (mostly subg. Guembelia) 14. Margins recurved on one or both sides. 15.
Leaves muticous, cucullate .......…17.
Grimmia atrata 15. Leaves awned, not cucullate. 16.
Leaves spirally curved around stem when dry, flagelliform innovations present
………………........…....…
30. Grimmia funalis 16.
Leaves straight when dry, 17.
Sporophytes present. 18.
Capsule exserted. 19.
Seta arcuate; dioicous. ..............
29. Grimmia elatior 19.
Seta straight; autoicous. 20.
Annulus prominent, 2 rows of rectangular cells; stomates in 2--3 rows; basal
juxtacostal cells sinuose and thick walled ...........14. Grimmia
longirostris 20.
Annulus small, 1 row of quadrate cells; stomates in 1 row; basal juxtacostal
cells straight and thin-walled ...........10. Grimmia
sessitana 18.
Capsule immersed. 21. Stem central strand present,
epidermis thin; leaf margins 2-stratose, not thickened, one margin recurved
(rarely both); leaves broadly ovate-lanceolate ...........
15. Grimmia arizonae 21. 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 .............
16. Grimmia pilifera 17.
Sporophytes absent. 22.
Basal juxtacostal cells straight, thin-walled; distal juxtacostal lamina
1-stratose, cells often bulging; plants small (< 1cm), blackish; moist,
alpine habitats ..........…..…..
10. Grimmia sessitana 22.
Basal juxtacostal cells sinuose, thick-walled; distal juxtacostal lamina at
least 2-stratose, cells not bulging; plants robust (> 1cm), yellow-green
to very dark olive green; dry, widely distributed. 23.
Leaf margins 2-stratose, not thickened; stem central strand present,
epidermis thin. 24.
Autoicous; costa transverse section reniform; leaves sheathing; widely
distributed .........14.
Grimmia longirostris 24.
Dioicous; costa transverse section semicircular; leaves not sheathing;
American Southwest ….........
15. Grimmia arizonae 23.
Leaf margins multistratose and thickened; stem central strand absent,
epidermis thick. 25. Leaves narrowly lanceolate from an
ovate base, usually narrowly recurved on both margins; distal lamina lacking multistratose bands, never
papillose .
16. Grimmia pilifera 25.
Leaves broadly lanceolate, broadly recurved on one margin; distal lamina with
multistratose bands, occasionally papillose ..
29. Grimmia elatior 14. Margins plane or incurved. 26.
Costa transverse section circular distally; awn hyaline-tipped to short (0.3
mm), decurrent . . . 13. Grimmia teretinervis 26.
Costa transverse section semicircular distally; awn short to long, usually
prominent, decurrent or not. 27.
Seta arcuate; leaves spirally curved around stem when
dry, flagelliform innovations present ….…………..…..…….....…… 30. Grimmia
funalis 27.
Seta straight; leaves straight when dry, flagelliform innovations absent. 28.
Gemmae abundant on adaxial distal leaf surface ..…...….……... 43. Grimmia
shastae 28.
Gemmae absent on leaves. 29.
Distal laminal cells not bulging. 30.
Margins plane throughout; basal marginal laminal cells short- to
long-rectangular; stomates present; autoicous. 31. 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 ........... 8. Grimmia donniana 31. 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 ….…...
10. Grimmia sessitana 30. Margins incurved distally; basal
marginal laminal cells quadrate to short-rectangular; stomates absent;
dioicous. 32. Capsule immersed to emergent,
wide-mouthed, peristome rudimentary; endemic to ..............................…
7. Grimmia mariniana 32. Capsule exserted, narrow-mouthed,
peristome present, fully developed; widespread in western 33. Basal juxtacostal laminal cells
short- to long-rectangular, distinct from quadrate to short-rectangular basal
marginal cells; medial laminal cells rounded, thick-walled.
........................ 5. Grimmia 33.
Basal juxtacostal and marginal cells quadrate to short-rectangular, basal
areolation relatively uniform; medial laminal cells quadrate to
short-rectangular, thin-walled .........................6. Grimmia
alpestris 29.
Distal laminal cells bulging. 34.
Leaf margins plane; cladautoicous; basal juxtacostal laminal cells
rectangular to elongate; distal juxtacostal laminal cells 1-stratose;
stomates present. …….
10. Grimmia sessitana 34.
Leaf margins incurved; dioicous; basal juxtacostal laminal cells quadrate to
short-rectangular; distal juxtacostal laminal cells 2-stratose; stomates
present or absent. 35.
Stomates present; leaves weakly to rarely strongly plicate distally,
cucullate …….… 11. Grimmia caespiticia 35. Stomates absent; leaves not
plicate, not cucullate ………. 6. Grimmia alpestris 6. Leaves merely 1-stratose distally; margins at most narrowly
2-stratose; one or both leaf margins recurved (G. reflexidens plane) .............…………………………...……… (mostly subg. Rhabdogrimmia) 36.
Costa projecting on dorsal side with two wings ....................................................................
39. Grimmia ramondii 36. Costa on dorsal side smooth or
angled. 37. Leaves muticous, without awns or only distal leaves with very short awn or hyaline tip. 38. Leaves cucullate ………….. 17. Grimmia atrata 38. Leaves not cucullate. 39.
Leaves squarrose when moist; medial laminal cells oblate to rounded quadrate
….…………..................... 34. Grimmia lisae 39.
Leaves erect to patent when moist; medial laminal cells quadrate to
short-rectangular. 40.
Basal marginal cells with thickened transverse walls; seta arcuate; globular
gemmae occasionally present on leaves .……..
.... 41. Grimmia trichophylla 40.
Basal marginal cells with thin transverse walls; seta straight or slightly
curved; gemmae absent 41.
Leaf margins both recurved; medial laminal cells with nodulose walls; seta
straight ............
25. Grimmia lesherae 41.
One leaf margin recurved; medial laminal cells sinuose; seta straight or slightly
curved ...……
9. Grimmia elongata 37. At least distal or perichaetial leaves with distinct awns. 42.
Awns nearly equal to or much longer than lamina, decurrent, seta straight to
flexuose. 43.
Awns 1--2 mm, leaf margins plane, [seta straight]; only known from ………...…………
12. Grimmia reflexidens 43.
Awns 2--4 mm, leaf margins both recurved; seta flexuose; endemic to 42. Awns typically
shorter than lamina (G. pulvinata
can be long); decurrent or not; seta arcuate to cygneous. 44.
Leaves abruptly narrowed to awns, apex rounded; autoicous. 45.
Operculum rostrate; basal marginal laminal cells quadrate .…………..... 38. Grimmia
pulvinata 45.
Operculum mammillate to conical; basal marginal laminal cells short- to
long-rectangular. 46.
Awns only on perichaetial leaves; basal juxtacostal cells with thin, straight
walls; marginal distal lamina 2-stratose; acidic rocks .. 35. Grimmia
moxleyi 46.
Awns on all distal leaves; basal juxtacostal cells with thick, nodulose
walls; marginal distal lamina 1-stratose; basic rocks .…..
37. Grimmia orbicularis 44.
Leaves gradually tapering to awns, apex narrowed; dioicous. 47. Clusters of gemmae easily visible
on dorsal side of leaf lamina or on leaf tips. 48. Multicellalar gemmae on dorsal side
of leaf lamina in distal leaves; leaves crisped and contored …..….
40. Grimmia torquata 48. Cluster of gemmae only on leaf
tips; leaves straight. 49. Leaves with blunt eroded apices;
distal cells with pseudo-papillose walls; stem central strand present ..............
26. Grimmia anomala 49. Leaves with acuminate apices;
distal cells with smooth walls; stem central strand absent ……….. 31. Grimmia hartmanii 47.
Clusters of gemmae absent or in obscure clusters in leaf axils. 50.
Leaves narrowly ovate-lanceolate to linear-lanceolate, distinctly incurved or
contorted when dry ..... 32. Grimmia incurva 50.
Leaves lanceolate to ovate-lanceolate, straight, appressed, only occasionally
somewhat contorted when dry. [51.
Shifted to left] 51. 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. 52.
Costa semicircular, not projecting in transverse section; basal marginal
cells thick-walled; distal leaves often secund, awns often decurrent
..................................................... 33. Grimmia leibergii 52.
Costa circular, projecting in transverse section; basal marginal cells
thin-walled; distal leaves straight, awns not decurrent
.………….........................................…….... 27. Grimmia attenuata 51. 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. 53.
Leaves squarrose when moist; mid-leaf cells rounded to oblate, straight
.……………........……… 34. Grimmia lisae 53.
Leaves erect to patent when moist; mid-leaf cells rounded-quadrate to
short-rectangular, sinuose. 54.
Yellowish green tufts; costa semicircular in transverse section; awns smooth
to slightly denticulate, not decurrent; capsule oblong-ovoid, yellow-green
………………....................…… 41. Grimmia trichophylla 54.
Blackish green tufts; costa angled or bluntly winged in transverse section;
awns denticulate, occasionally decurrent; capsule globose, brown
………………………….......... 36. Grimmia muehlenbeckii xxxa. Grimmia
subg. Gasterogrimmia Schimper, Coroll. 46. 1856 Plants to 15 mm. Stem
central strand strong. Stem leaves
oblong-ovate to oblong-lanceolate, keeled or concave, margins plane, costa
prominent, distal lamina 2-stratose (1-stratose for G. plagiopodia); juxtacostal and marginal basal laminal cells
hyaline. Gemmae absent. Seta arcuate to sigmoid (except G. Species 4 (4 in the flora): North America, Mexico,
Central America, South America, Eurasia, n. Africa, Pacific Islands (New
Zealand), Australasia, Antarctic. Members of this subgenus are recognized by their
commonly occurring immersed, ventricose capsules on arcuate to sigmoid setae
that are eccentrically attached. Additionally, all these species have three
to four large stomates at the very base of the capsule. The entire basal
region of the leaf is typically hyaline. Most members of the subgenus are
restricted to calcareous habitats. The genus Jaffueliobryum is the only other group in the Grimmiaceae to be
so strongly tied to calcareous substrates. 1. Grimmia plagiopodia
Hedwig, Spec. Musc. 78. 15 f. 1801 Grimmia brandegei Plants in dense cushions to hoary tufts, dark green to
brown. Stems 0.3--0.5(--1) cm, Stem
leaves oblong-ovate, 1--1.7 × 0.4--0.8 mm, concave-keeled, awn 0.3--1 mm;
distal laminal cells 1-stratose, marginal cells 1-2-stratose; medial laminal
cells quadrate to short-rectangular, slightly sinuose, slightly thick-walled;
basal juxtacostal laminal cells quadrate to short-rectangular, straight,
thin-walled; basal marginal laminal cells quadrate to short-rectangular,
straight, thin-walled. Sexual
condition gonioautoicous. Seta
sigmoid, 0.2--0.3 mm. Capsule
usually present, exothecial cells thin-walled, annulus absent, operculum
mammillate, peristome present, fully developed, perforated and split in
distal half. Exposed calcareous sandstone, limestone,
occasionally concrete, and glacio-lacustrine silt; 50--2400 m; Greenland;
Alta., B.C., N.W.T., Nunavut, Ont., Sask.; Alaska, Calif., Col., Idaho, Ill.,
Iowa, Minn., Mont., Neb., Nev., N.Mex., N.Dak., S.Dak., Utah, Wis., Wyo.;
South America; Eurasia; Pacific Islands (New Zealand); Antarctic. Grimmia plagiopodia has a widespread and continuous distribution on
calcareous rock across the northern Great Plains of North America, reaching
as far east as 2. Grimmia anodon Bruch
& Schimper, Bryol. Euro. 3: 110. 1845 Grimmia anodon var. anomala
Bartram; G. subanodon Ochyra; Schistidium obtusifolium Ireland &
H. A. Crum Plants in small cushions, dark green to brown. Stems
0.5--1 cm. Stem leaves
oblong-ovate to oblong-lanceolate, 0.9--2 × 0.4--0.8 mm, concave-keeled, awn
0.1--1.2 mm; distal laminal cells 1-stratose with 2-stratose patches,
marginal cells 2-stratose; medial laminal cells quadrate, sinuose,
thick-walled; basal juxtacostal laminal cells quadrate to long-rectangular,
straight, thin-walled; basal marginal laminal cells quadrate to
long-rectangular, straight, thin-walled. Sexual
condition gonioautoicous. Seta
sigmoid, 0.2--0.3 mm. Capsule
usually present, exothecial cells thin-walled, annulus of 1--2 rows of
quadrate, thin-walled cells, not revoluble, operculum mammillate, peristome
absent. Exposed, calcareous sandstone, limestone, and concrete; 25--2700 m; Greenland; Alta, B.C., N.B., N.W.T., Nunavut, Ont., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Idaho, Mich., Mont., Nev., N.Mex., N.Y., N.Dak., Oreg., S.Dak. Utah, Wash., Wyo.; ne Mexico; South America (Bolivia, Chile); Eurasia; Africa (Morocco). Grimmia anodon is widespread and common across the western 3. Grimmia americana Bartram,
Bryologist 32: 8. 1929 Plants in hoary tufts, dark green to brown. Stems 0.5--1.5 cm. Stem leaves oblong-ovate to
oblong-lanceolate, 1.5--2 × 0.5--0.9 mm, concave, awn to 1 mm; distal laminal
cells 2-stratose, marginal cells 2-stratose; medial laminal cells
rounded-quadrate, slightly thick-walled; basal juxtacostal laminal cells
quadrate to short-rectangular, straight, thick-walled; basal marginal laminal
cells quadrate to long-rectangular, straight, thick-walled. Sexual condition gonioautoicous. Seta straight to arcuate, 0.6--1 mm. Capsule usually present, exothecial
cells thick-walled, annulus of 2--3 rows,
of rectangular, thin-walled cells, revoluble, operculum rostellate,
peristome present, fully developed, perforated and split in most distal part,
weakly papillose. Calcareous rock; of
conservation concern; around 1600 m; Grimmia 4. Grimmia poecilostoma Cardot
et Sebille, Rev. Bryol. 28: 118. 1901 Grimmia tergestina var. poecilostoma
Loeske Plants in loose tufts, olive green to black. Stems 0.5--0.9 cm. Stem leaves oblong-ovate to
oblong-lanceolate, 1.6--2 × 0.3--0.6 mm, concave, awn 0.3--0.6 mm; distal
laminal cells 2-stratose, marginal cells 2-stratose; medial laminal cells
rounded, straight, thick-walled; basal juxtacostal laminal cells quadrate to
long-rectangular, straight, thin-walled; basal marginal laminal cells
quadrate to short-rectangular, straight, thin-walled. Sexual condition dioicous. Seta
sigmoid, 0.3--0.5 mm. Capsule
usually present, exothecial cells thin-walled, annulus of 2--3 rows,
rectangular, thick-walled, revoluble, operculum obliquely rostrate, peristome
present, rudimentary, perforated, papillose. Basalt, granite, schist and
limestone; 500--2100 m; B.C., N.W.T., In North
America, Grimmia poecilostoma is
known from only scattered localities in the American west and in three
extremely disjunct sites in xxxb. Grimmia
subg. Guembelia (Hampe) Schimper,
Coroll. 48. 1856 Guembelia Hampe, Bot. Zeitung 4: 124. 1846 Plants 10--20 mm (--70 mm for G. atrata). Stem
central strand present or absent. Stem
leaves narrowly ovate-lanceolate to oblong-lanceolate, keeled, margins
plane, recurved or incurved, costa prominent, distal lamina 2-stratose or
with at least several marginal rows 2-stratose; basal marginal cells hyaline
or not. Gemmae absent. Sexual condition autoicous or
dioicous. Seta straight, centrally
attached to capsule. Capsule
immersed to exserted, smooth (except G.
mariniana); stomates present or absent, small, at neck to part way up
capsule; calyptra mitrate to cucullate, covering operculum. Species 41 (13 in the flora): North America, Members of this variable subgenus are recognized by
their thick, keeled leaves and usually long, straight setae with smooth
capsules. Most members of the subgenus occur on dry acidic rock, with three
species growing on damp to wet acidic rock and one on dry calcareous rock. 5. Grimmia Grimmia arctophila Kindberg; G.
brachydon Austin; G. jamesii
Austin; G. montana var. brachydon (Austin) Lesquereux and
James; G. tenella (J. K. A. Müller)
Kindberg; Guembellia tenella J.K.A.
Müller Plants in hoary cushions, yellow-green to dark blue-green,
sometimes almost black. Stems 1--1.2(--1.5)
cm, central strand weak. Stem leaves
narrowly lanceolate, rarely ovate-lanceolate, 1--2 × 0.3--0.6 mm,
concave-keeled, not plicate, margins plane, usually narrowly incurved
distally, awn 0.2--1.3 mm, costal transverse section not prominent to
prominent, semicircular; distal laminal cells 2-stratose, not bulging,
marginal cells 2-stratose, not bulging; medial laminal cells rounded,
thick-walled; basal juxtacostal laminal cells short- to long-rectangular,
straight, thick-walled; basal marginal laminal cells quadrate to
short-rectangular, straight, thick-walled, not hyaline. Perichaetial leaves not enlarged. Sexual condition dioicous. Seta
straight, 2--3 mm. Capsule
occasionally present, exserted, yellow to brown, oblong, exothecial cells
rectangular, thin-walled, stomates absent, annulus of 1 row of quadrate,
thick-walled cells, operculum rostellate, peristome present, fully developed,
split and perforated in distal half. Exposed acidic granite and sandstone; 900--4000 m; Greenland; Alta., B.C., Nunavut, Ont., Yukon; Alaska, Ariz., Calif., Col., Idaho, Mont., Nev., N.Mex., Oreg., Utah, Wash., Wyo.; Mexico; Europe; Africa. Grimmia 6. Grimmia alpestris (Weber
& Mohr) Schleicher, Cat. Pl. Helv. Ed. 2: 29. 1808 Grimmia donniana var. alpestris
(Weber & Mohr) Hampe; Trichostomum
pulvinatum var. alpestre Weber
& Mohr, Bot. Taschenbuch: 110. 1807 Plants in cushions or mats, yellow-green, glaucous-green
to dark green, sometimes almost black. Stems
1--1.2(--1.5) cm, central strand weak. Stem leaves narrowly lanceolate to ovate-lanceolate, 1--1.8 ×
0.2--0.6 mm, keeled, not plicate, margins plane proximally, incurved
distally, awn 0.3--0.8 mm, costal transverse section prominent, semicircular;
distal laminal cells 2-stratose, bulging or not, marginal cells 2-stratose;
medial laminal cells quadrate to short-rectangular, straight, thin-walled;
basal juxtacostal laminal cells quadrate to short-rectangular, straight,
thick-walled; basal marginal laminal cells quadrate to short-rectangular,
straight, thick-walled, 1--2 rows hyaline or not hyaline. Perichaetial leaves not enlarged. Sexual condition dioicous. Seta straight, 2--3 mm. Capsule occasionally present,
exserted, yellow to brown, ovate to oblong-ovate, exothecial cells quadrate
to short-rectangular, thick-walled, stomates absent, annulus of 1 row of
quadrate, thick-walled cells, operculum conic to mammilate with a short
obtuse beak, peristome present, fully developed, split and perforated in
distal half. Exposed acidic granite and sandstone; 360--3300 m;
Greenland; Alta., B.C.; Ariz., Calif., Col., Idaho, Mont., N.Mex., Nev.,
Oreg., S. Dak., Utah, Wash., Wyo.; Eurasia. Grimmia alpestris has a distribution similar to that of G. 7. Grimmia mariniana Sayre,
Bryologist 58: 323. 1955 Plants in hoary loose cushions, emerald green to black. Stems 0.6--1.2 cm, central strand
weak. Stem leaves
linear-lanceolate to narrowly ovate-lanceolate, 1.5--2.1 × 0.5--0.7 mm,
keeled, not plicate, margins incurved distally, awn 0.4--0.8 mm, costal
transverse section prominent, semicircular; distal laminal cells 2-stratose,
not bulging, marginal cells 2-stratose, not bulging; medial laminal cells
quadrate, sinuose, thick-walled; basal juxtacostal laminal cells
short-rectangular to elongate, sinuose, thick-walled; basal marginal laminal
cells quadrate to short-rectangular, straight, thick-walled, hyaline. Perichaetial leaves slightly
enlarged. Sexual condition
dioicous. Seta straight, to 1 mm. Capsule occasionally present,
immersed to emergent, yellow (rarely brown), ovate with open mouth, faintly
but distinctly ribbed, exothecial cells rectangular, thick-walled, stomates
absent, annulus of 1 row of quadrate, thick-walled cells, operculum
rostellate, with a short obtuse beak, peristome present, but rudimentary,
teeth composed of only a few basal cells, sometimes perforated. Acidic sedimentary and basaltic metavolcanics; of conservation
concern; 900--1200 m; Grimmia mariniana is a rare endemic, known only from the coastal
mountains of central 8. Grimmia donniana Smith, Engl. Bot. 18. 1259. 1804 Plants in dense cushions, dark green to almost black. Stems 0.8--1.2(--1.5) cm, central
strand present. Stem leaves
oblong-lanceolate, 1--2.2 × 0.3--0.6 mm, keeled, not plicate, margins plane,
awn 0.3--1.3 mm, costal transverse section prominent, semicircular; distal
laminal cells commonly 2-stratose, occasionally only 1-stratose, not bulging,
marginal cells 2-stratose, not bulging; medial laminal cells
short-rectangular, sinuose, thick-walled; basal juxtacostal laminal cells
long-rectangular, straight, thin-walled (rarely somewhat thick-walled); basal
marginal laminal cells long-rectangular, straight, thin-walled, typically
hyaline. Perichaetial leaves not
enlarged. Sexual condition
autoicous. Seta straight, 2--3 mm.
Capsule usually abundantly
present, exserted, pale yellow-brown, oblong, exothecial cells quadrate,
thin-walled, stomates present, annulus of 2 rows of quadrate, thick-walled
cells, operculum mammillate to rostellate, peristome present, fully
developed, perforated in distal half. Exposed, acidic granite and sandstone, high
elevation forests to tundra; 880--3700 m; Greenland; Alta., B.C., Labr.,
N.W.T., Que., Yukon; Alaska, Colo., Idaho, Maine, Mich., Mont., N.H., N.Y.,
Oreg., Utah, Wash., Wyo.; Mexico; South America; Eurasia; Africa; Antarctic. Grimmia donniana is widespread but relatively uncommon and sporadic
along the front ranges of the Rocky Mountains from 9. Grimmia elongata Kaulfuss in
J. W. Sturm, Deutschl. Fl. Abth. II, Crypt. 4(13): 24.
1812 Drypotodon elongatus (Kaulfuss) Hartman Plants in dense
patches, reddish brown to blackish green. Stems 1--4 cm, central
strand present. Stem leaves lanceolate, keeled, not plicate, one
margin commonly narrowly recurved, awns 0.1--0.5 mm, lower leaves muticous,
costal transverse section weak at base, semicircular distally; distal laminal cells 1-stratose, not
bulging, marginal cells 2-stratose, not bulging; medial laminal cells short-rectangular, sinuose, thick-walled;
basal juxtacostal laminal cells
elongate, straight, somewhat thick-walled; basal marginal laminal cells
rectangular with straight walls, often hyaline. Perichaetial leaves not enlarged. Sexual condition dioicous. Seta
straight to slightly curved, 1.5--2.5 mm. Capsule occasionally present, emergent
to shortly exserted, yellow-brown, ovoid, exothecial cells variable, quadrate
to rectangular, thin-walled, stomates present, annulus of 1--3 rows,
operculum conical to rostrate, peristome present, fully developed, papillose. Damp acidic volcanic
rock and sandstone, high elevation forests to tundra; 400--2400 m; Greenland;
Alta., N.W.T., Nunavut, Que.; Colo.; Mexico; Central America; South America;
Eurasia; Africa. Grimmia
elongata is a
widely distributed species mainly occurring above 2000 m, and with a
preference for acidic sandstones and volcanic, damp, north-facing outcrops
and ledges. It seems to be uncommon in 10. Grimmia sessitana De
Notaris, Atti Reale Univ. Genova 1: 704. 1869 Grimmia subpapillinervis Kindberg; G. tenerrima Renauld & Cardot Plants in dense patches, dark green, brown-green,
sometimes almost black. Stems
0.5--1 cm, central strand strong. Stem
leaves lanceolate to ovate-lanceolate, 1--1.8 × 0.2--0.6 mm, keeled, not
plicate, one margin commonly recurved, sometimes both plane, awn 0.3--0.8 mm,
costal transverse section prominent, semicircular; distal laminal cells
1-stratose, often bulging, marginal cells widely 2-stratose, often bulging;
medial laminal cells quadrate, sinuose, thick-walled; basal juxtacostal
laminal cells rectangular to elongate, straight, thin-walled; basal marginal
laminal cells short- to long-rectangular, straight, thick transverse and thin
lateral walls, hyaline or not.
Perichaetial leaves not enlarged. Sexual
condition cladautoicous. Seta
straight, 1--2.5 mm. Capsule
usually present, exserted, yellow, ovate to elliptic, exothecial cells
short-rectangular, thin-walled, stomates present, annulus of 1 row of
quadrate, thick-walled cells, operculum conic, usually with a short obtuse
beak, peristome present, fully developed, split or solid in distal half. Exposed or sheltered, moist, acidic granite and sandstone,
alpine; 1100--3900 m; Greenland; Alta., B.C., Labr., Nfld., N.W.T., Quebec,
Yukon; Alaska, Ariz., Calif., Colo., Idaho, Mont., N.H., N.M., N.Y., Nev.,
Oreg, Utah, Wash., Wyo.; South America (Argentina, Chile); Eurasia; Africa
(Kenya, South Africa, Uganda); Antarctic. Grimmia sessitana is widely distributed in high elevation sites
throughout western North America, principally in mountains forming the
continental divide, but also in the northern Sierra Nevada and the coastal
range of 11. Grimmia caespiticia (Bridel)
Juratzka. Laubm.-Fl. Oesterr.-Ung.: 172. 1882 Campylopus caespiticius Bridel, Muscol. Recent. Suppl. 4: 77; Grimmia alpestris var. caespiticia (Bridel) D. N. Jones; G. alpestris var. manniae (C. Müller) D. N. Jones; G. alpestris var. holzingeri
(Cardot & Theriot) D. N. Jones; G.
manniae C. Müller; G. nivalis
Kindberg; G. pyrenaica Kern Plants in dense flat mats, blue-green to black-green. Stems
0.5--1 cm, central strand weak. Stem
leaves lanceolate from a broad base, 0.8--1.3 × 0.2--0.5 mm, keeled,
weakly to rarely strongly plicate distally [often strongly plicate in
Eurasian specimens], margins plane proximally, incurved distally, cucullate,
awn 0.1--0.5 mm, often muticous, costal transverse section prominent,
semicircular; distal laminal cells 2-stratose, bulging, marginal cells 2-stratose,
bulging; medial laminal cells rounded-quadrate, thick-walled; basal
juxtacostal laminal cells quadrate to short-rectangular, straight,
thin-walled; basal marginal laminal cells short- to long-rectangular,
straight, thick transverse and thin lateral walls, not hyaline. Perichaetial leaves not enlarged. Sexual condition dioicous. Seta straight, 1.8--2.4 mm. Capsule occasionally present,
exserted, yellow, cylindric, exothecial cells short-rectangular, thin-walled,
stomates present, annulus of 1 row of quadrate, thick-walled cells, operculum
mammillate, peristome present, fully developed, solid in distal half. Exposed, dry to moist, acidic granite and quartzite,
alpine; 1200--3500 m; Greenland; Alta., B.C.; Grimmia caespiticia is an uncommon species that occurs on siliceous
rock outcrops above timberline in western Grimmia caespiticia may also be confused with G. sessitana. These species are both found above timberline and
both have bulging laminal cells and capsules with stomates. However, the
incurved leaf margins, cucullate apex, and quadrate to short-rectangular
basal areolation of G. caespiticia
is quite different from the plane to recurved leaf margins with
long-rectangular basal areolation typical of G. sessitana. Although the type specimen of G. alpestris var. holzingeri
lacks capsules, gametophytically it is indistinguishable from muticous
specimens of G. caespiticia.
Specimens of var. holzingeri with
capsules have been collected near the type locality and these specimens have
stomates. Rather than accepting that G.
alpestris may have stomates (in the sense of 12. Grimmia reflexidens C.
Müller., Syn. Musc. Frond. 1: 795. 1849 Grimmia grisea Cardot Plants in compact hairy cushions, greyish
green. Stems 1--2 cm, central
strand absent. Stem leaves ovate to oblong-lanceolate, 0.1--1.15 ×
0.3--0.5 mm, keeled, not plicate, margins plane, awns 1--2 mm, very long,
smooth to slightly denticulate, flattened basally, long-decurrent, costal
transverse section prominent, semicircular; distal laminal cells yellowish-green, 1-stratose with 2-stratose
ridges, not bulging, marginal cells 2-stratose, not bulging; medial laminal cells
rounded-quadrate, slightly sinuose, thick-walled; basal juxtacostal laminal cells
rectangular, sometimes nodulose, thin- to thick-walled; basal marginal laminal cells rectangular
with thickened transverse walls, pellucid in 2--4 rows. Perichaetial leaves enlarged. Sexual condition dioicous. [Seta
straight, 1.5--2 mm. Capsule absent in northern hemisphere
material, emergent to shortly exserted, yellowish brown, oblate, exothecial
cells irregularly short-rectangular, thin-walled, stomates absent, annulus of
1 row of quadrate, thick-walled cells, operculum mammillate to rostrate, peristome
present, nearly fully developed, split and perforated only in apex,
papillose.] Dry
acidic rock; 50 [--300] m; of conservation concern; Nfld.; South America
(Argentina, Chile); Europe (Iceland); Australia; Pacific Islands (New
Zealand). Previous to its discovery in 13. Grimmia teretinervis Limpricht, Jahresber. Schles. Ges. Vaterl.
Cult. 61: 216. 1884 Schistidium teretinerve (Limpricht) Limpricht Plants in loose tufts, green-brown to reddish brown,
shiny. Stems 2--3 cm, central
strand strong. Stem leaves
ovate-cordate to lanceolate, 0.6--1.2 × 0.2--0.5 mm, keeled, not plicate,
margins plane, awn to 0.3 mm, often just hyaline tipped, commonly
long-decurrent, costal transverse section prominent, circular distally;
distal laminal cells 2-stratose, bulging, marginal cells 2-stratose, bulging;
medial laminal cells rounded-quadrate, thick-walled; basal juxtacostal
laminal cells quadrate to short-rectangular, straight, thin- to thick-walled;
basal marginal laminal cells oblate to quadrate, straight, thick-walled, not
hyaline. Perichaetial leaves
unknown. Sexual condition
dioicous. Seta unknown. Capsule unknown. Moist calcareous sandstone, limestone and dolomite
outcrops; 250--1700 m; Alta., B.C., N.W.T., Ont., Que., Sask., Yukon; Alaska,
Ark., Colo., Minn., Mo., Mont., Nev., Okla., Wis.; Europe. As reported by R. R. Ireland (1982), Grimmia teretinervis is widely
scattered across 14. Grimmia longirostris Hooker,
Musci Exot.: 62. 1818 Grimmia affinis Hornschuch; G.
arctophila subsp. labradorica
Kindberg; G. catalinensis Bartram; G. catalinensis var. mutica Bartram; G. elata Kindberg.; G.
ortholoma Kindberg; G. ovalis
var. affinis (Hornschuch)
Brotherus; G. ovata var. affinis (Hornschuch) Bruch &
Schimper; G. ovateoformis Kindberg Plants in compact cushions, yellow-green to dark olive
green. Stems 1--3 cm, central
strand strong. Stem leaves
ovate-lanceolate, 1.5--3 × 0.6--0.7 mm, keeled, one margin recurved
proximally, not sheathing, awn 0.5--1.5 mm, costal transverse section prominent,
reniform; distal laminal cells 2-stratose, not bulging, marginal cells
2-stratose, not bulging; medial laminal cells short-rectangular, sinuose,
thick-walled; basal juxtacostal laminal cells long-rectangular to linear,
sinuose, thick-walled; basal marginal laminal cells short-rectangular,
straight, thick transverse and thin lateral walls, hyaline. Perichaetial leaves not enlarged. Sexual condition cladautoicous. Seta straight, (1--)2--4 mm. Capsule usually present, (emergent
to) exserted, yellow, oblong-ovoid to cylindric, exothecial cells short- to
long-rectangular, thin-walled, stomates present in 2--3 rows, annulus of 2
rows of rectangular, thick-walled cells, operculum long-rostrate, peristome
present, fully developed, split and perforate in distal half. Exposed, dry, acidic granite and quartzite;
100--3050 m; Greenland; Alta., B.C., Labr., Man., N.B., Nfld., N.W.T, N.S.,
Nunavut, Ont., Que., Sask., Yukon; Alaska, Ariz., Calif., Colo., Idaho,
Maine, Minn., Mont., Nev., N.H., N.Mex., N.C., Okla., Oreg, S.Dak., Tex.,
Utah, Vt., Wash., Wyo.; Mexico; Central America (Guatemala, Honduras, Costa
Rica); South America; Eurasia; Africa; Pacific Islands; Australia. Grimmia longirostris is one of the most common species of Grimmia. It is most common in the
eastern ranges of the Rocky Mountains, ranging from western As Grimmia
affinis Hornschuch, G. longirostris
has commonly been placed as a subspecific entity of Grimmia ovalis. Despite G. Sayre’s (1951) resolution of the
differences between these taxa, a large proportion of specimens in major
herbaria in 15. Grimmia arizonae Renauld
& Cardot, Rev. Bryol. 19: 85. 1892 Grimmia santa-ritae Bartram Plants in hoary tufts, olive green to dark blue-green. Stems 1--3 cm, central strand strong.
Stem leaves ovate-lanceolate,
1.5--3 × 0.6--0.7 mm, keeled, one margin recurved proximally (occasionally
both), sheathing, awn 0.5--1.5 mm, costal transverse section prominent,
reniform to semicircular; distal laminal cells 2-stratose, not bulging,
marginal cells 2-stratose, not bulging; medial laminal cells
short-rectangular, sinuose, thick-walled; basal juxtacostal laminal cells
long-rectangular to linear, sinuose, thick-walled; basal marginal laminal
cells short-rectangular, straight, thick transverse and thin lateral walls,
hyaline. Perichaetial leaves not
enlarged. Sexual condition
dioicous. Seta straight, 0.5--0.7
mm. Capsule occasionally present,
immersed, yellow, oblong-ovoid, exothecial cells quadrate, thin-walled,
stomates present in 2--3 rows, annulus of 2 rows of rectangular, thick-walled
cells, operculum long-rostrate, peristome present, perforate in middle, split
in distal half. Exposed, dry, basalt and acidic granite, rarely on
sandstone; 1800--2700 m; Grimmia arizonae is endemic to the American Southwest and northern As discussed above, Grimmia arizonae is part of a group that includes G. longirostris and G. pilifera. Its sheathing leaf bases,
dioicous sexuality and immersed capsules will separate it from G. longirostris. The separation of Grimmia arizonae from Grimmia
pilifera, however, has often proven problematic. Grimmia pilifera has been described as having strongly keeled
leaves and margins 2--3-stratose, thicker than the medial lamina. (e.g. H. A.
Crum 1994). H. C. Greven (1999) adds with usually with short awned,
ovate-lanceolate leaves. Grimmia
arizonae has less keeled leaves and margins 2-stratose, not thicker than
lamina. Greven notes its usually long awns and broadly lanceolate
leaves. J. Muñoz (1999a),
however, synonymized these species, attributing any differences as due to
geographical variations and sexual development. He reports that in eastern 16. Grimmia pilifera P.
Beauvois, Prodr. Aetheogam: 58. 1805 Grimmia pensylvanica Schwaegrichen Plants in robust, loose tufts, dark olive green to black. Stems 1--4 cm, central strand absent.
Stem leaves narrowly lanceolate
from an ovate base, 2--4.5 × 0.4--0.8 mm, keeled, both margins recurved
proximally, often narrowly, sheathing, awn 0.2--0.6(--1.5) mm, costal
transverse section prominent, usually semicircular; distal laminal cells
2-stratose, not bulging, marginal cells (2--)3(--4)-stratose, not bulging;
medial laminal cells quadrate to short-rectangular, sinuose, thick-walled;
basal juxtacostal laminal cells short-rectangular to linear, sinuose, thin
transverse and thick lateral walls; basal marginal laminal cells quadrate to
short-rectangular, straight to sinuose, thick transverse and thin lateral
walls, hyaline. Perichaetial leaves
not enlarged. Sexual condition
dioicous. Seta straight, 0.5--1
mm. Capsule occasionally present,
immersed, yellow, oblong-ovoid, exothecial cells quadrate to
short-rectangular, thin-walled, stomates present in 2--3 rows, annulus of 2
rows of rectangular, thick-walled cells, operculum long-rostrate, peristome
present, fully developed, perforate in middle, split in distal half. Exposed to tree shaded, dry limestone, sandstone and
granite; 60--2300 m; N.S.; Ala., Ariz., Ark., Colo., Conn., Del., Ga., Ill.,
Ind., Ky., Maine, Md., Mass., Minn., Mo., Mont., N.J., N.Mex., N.Y., N.C.,
Pa., R.I., S.Dak., Tenn., Vt., Va., W.Va.; Mexico; Asia. Grimmia
pilifera is most
commonly found in the Appalachian Mountains from 17. Grimmia atrata Hornschuch,
Flora 2: 85. 1819 Dryptodon atratus (Hornschuch) Hartman Plants in
variable loose patches, dark green to black, frequently rust colored below. Stems
2--7 cm, central strand absent. Stem leaves lanceolate to
ligulate, 1.5--3 × 0.3--0.6 mm, keeled, margins recurved proximally, incurved
distally, tapering to blunt cucullate apex, muticous, costal transverse section prominent, usually semicircular; distal laminal cells 1-stratose with
2-stratose ridges, to completely 2-stratose; medial laminal cells rectangular, nearly straight to
sinuose or nodulose, thick-walled; basal juxtacostal laminal cells
rectangular, straight to slightly sinuose walled; basal marginal laminal cells in 1--3 rows
quadrate, hyaline with straight to slightly sinuose, thick transverse walls. Perichaetial leaves not enlarged. Sexual condition dioicous. Seta straight,
2--6 mm. Capsule occasionally present, long-exserted, yellow-brown,
oblong to cylindric, exothecial cells rectangular, thick-walled, annulus of
3--4 rows of rectangular, thick-walled cells, stomates present, operculum
conic to rostrate, peristome present, fully developed, perforated and split
distally, weakly papillose. Damp, heavy-metal-bearing rock from the lowlands to
the alpine; of conservation concern; 1100--2600 m; Greenland; Labr., Yukon;
South America (Bolivia); Eurasia. Grimmia atrata is rare in xxxc. Grimmia subg. Litoneuron Hagen, K.
Norsk. Vid. Selsk. Skrift. 1909 (5): 6, 10. 1909 Plants 1--4 cm. Stem
central strand present. Stem
leaves broadly oblong-ovate to narrowly ovate-lanceolate or ligulate from
an ovate base, concave, margins plane or incurved, costa immersed, distal
lamina 2--3-stratose; basal marginal cells hyaline or not. Gemmae absent. Sexual condition dioicous. Seta
straight or sigmoid, centrally attached to capsule. Capsule exserted (emergent in G.
nevadensis), smooth; stomates present or absent, small, at base of
capsule; calyptra mitrate to cucullate, covering operculum. Species 15 (7 in flora): North America; Eurasia;
Africa; Mexico; Central America; South America; Indian Ocean Islands;
Australia. Members of this subgenus are
recognized by their thick, concave leaves with plane margins and costa not
projecting prominently from the lamina. The stems have well-developed central
strands. They are dioicous, and capsules are not common, but if present they
are usually long-exserted on a straight or rarely only slightly sigmoid seta.
The annulus is typically large and prominent but is lacking in Grimmia hamulosa. Members of the
subgenus have broad habitat preferences ranging from dry exposed rocks to
stream banks and cliff faces along lake shores. They are found on acidic to
basic rocks ranging from near sea level up to the alpine zone. 18. Grimmia unicolor Hooker
in R. K. Greville, Scott. Crypt. Fl. 3: 123. 1825 Plants in dense to loose patches, pale green to red-brown.
Stems 1.5--4(--5) cm. Stem leaves narrowly
oblong-lanceolate to ligulate from an ovate base, 1.5--2.5 × 0.5--0.7 mm,
both margins incurved, intermarginal bands absent, often sheathing, muticous,
cucullate, obtuse-rounded, costa narrow proximally; distal laminal cells
2--3-stratose, rounded, thick-walled; medial laminal cells rounded to
quadrate, straight, thick-walled; basal juxtacostal laminal cells
short-rectangular, straight, thick lateral walled, pale yellow; basal
marginal laminal cells short-rectangular, straight, thick lateral walled,
pale yellow, hyaline. Perichaetial
leaves enlarged. Seta straight
to slightly sigmoid, 2--4 mm. Capsule
occasionally present, exserted, brown, oblong-ovoid, exothecial cells
short-rectangular, thin-walled, stomates present, annulus of 2--3 rows of
rectangular, thick-walled cells, operculum long-rostrate, peristome perforate
and split in distal half. Calyptra
mitrate. Cracks of wet acidic, siliceous rocks especially along streams or splash zones of lake shores; 225--2000 m; Greenland; B.C., Ont., Que., Nfld.; Alaska, Calif., Maine, Mich., Minn., N.Y., Vt.; Eurasia; Africa (Ethiopia). In 19.
Grimmia hamulosa Lesquereux, Mem. California
Ac. Sc. 1: 14. 1868 Grimmia brevirostris
Williams Plants in flat
patches, blackish green. Stems
1--1.5 cm. Stem leaves homomallous-falcate, oblong-lanceolate, from a
clasping base tapering to a long, subulate uncinate point, 2--3.5 mm, margins
plane, erect above, intermarginal bands absent,
usually muticous, occasionally short hyaline awns are present, not
decurrent, costa narrow proximally; distal
laminal cells 2-stratose, irregularly rounded to quadrate; medial
laminal cells irregularly rounded to quadrate, sinuose, thick-walled; basal juxtacostal laminal cells
short-rectangular, straight to sinuose, thick-walled; basal marginal laminal cells quadrate to
short-rectangular, thick-walled. Perichaetial leaves not enlarged. Seta straight,
2.5--3.5 mm. Capsule occasionally present, exserted, oblate, shiny,
smooth, becoming striate when dry, stomates absent, annulus absent, operculum
conical, peristome split in distal half, basal segments smooth, distal
segments papillose. Calyptra unknown.
Dry
granitic rock and boulders; of conservation concern; 1500--3500 m; Grimmia
hamulosa is a rare endemic to the 20. Grimmia olneyi Sullivant
in W. S. Sullivant & L. Lesquereux, Musc.
Bor.-Amer.: 32. 1856 Grimmia austinii Kindberg Plants in flat patches, dark green to brownish black. Stems 1--2 cm. Stem leaves narrowly ovate-lanceolate from an ovate base, 2--3 ×
0.4--0.8 mm, both margins incurved, intermarginal bands absent, awn 0.1--0.5
mm, not decurrent, acute, costa narrow proximally; distal laminal cells 2--4
stratose, rounded, thick-walled; medial laminal cells quadrate, slightly
thick-walled; basal juxtacostal laminal cells quadrate to short-rectangular,
straight, thick lateral walls, green; basal marginal laminal cells quadrate,
straight, thick transverse and thin lateral walls, green, hyaline. Perichaetial leaves not enlarged. Seta sigmoid, 3--4 mm. Capsule occasionally present,
exserted, brown, oblong-ovoid, exothecial cells short-rectangular,
thin-walled, stomates present, annulus of 2--3 rows of rectangular,
thick-walled cells, operculum long-rostrate, peristome perforate in distal
half, split in distal half. Calyptra
cucullate. Cracks and exposed faces of
dry to periodically wet, acidic or calcareous rocks, commonly along streams
or splash zones of lake shores; 20--600 m; Ont., N.S.; Ark., Conn., Del.,
Maine, Mass., Mich., Mo., N.H., N.J., N.Y., N.C., Pa., R.I., Tenn., Vt., Va. Grimmia olneyi is endemic to eastern North America with its center
of distribution being along the Appalachians from the New England states
southward to Grimmia olneyi and G.
laevigata have broadly overlapping distributions in eastern 21. Grimmia ovalis (Hedwig)
Lindberg, Acta Soc. Sci. Fenn. 10: 75. 1871 Dicranum ovale Hedwig, Sp. Musc. Frond.: 140. 1801; Grimmia commutata Huebener; G. ovata Weber & Mohr; G. ovata var. gracilis Röll Plants in loose tufts, dark green to brownish black. Stems 1--3 cm. Stem 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; distal laminal cells 2-stratose, quadrate,
thick-walled; medial laminal cells rounded to quadrate, straight,
thick-walled; 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. Perichaetial leaves enlarged. Seta straight, 4--6 mm. Capsule occasionally present,
exserted, yellow-brown, oblong-ovoid, exothecial cells short-rectangular,
thin-walled, stomates present, annulus of 2--3 rows of rectangular,
thick-walled cells, operculum long-rostrate, peristome solid, split in distal
half. Calyptra cucullate. Dry, exposed to partially
shaded, acidic, sandstone and granite and basalt, montane to alpine;
(30--)1000--2450 m; Greenland; Alta, B.C., Ont., Que., Yukon; Ariz., Calif.,
Colo., Idaho, Minn., Mo., Mont., Nev., N.Mex., N.Y., Ore., Pa., Tex., Utah,
Va., Wash., Wyo.; Eurasia; Africa (Algeria, Morocco). Grimmia ovalis is common and widespread in high elevation sites in
western North America from southern B.C. along the Based on identifications by H. C. Greven, W. A.
Weber et.al. (2003) reported Grimmia
bernoullii in the 22. Grimmia nevadensis H. C.
Greven, Bryologist 105: 273. 2002 Plants in dense cushions, blackish green. Stems 0.5--0.7(--1) cm, central
strand weak. Stem leaves
ovate-lanceolate, 1.5--2 mm, both margins plane, incurved distally,
intermarginal bands absent, awn 0.4--0.8 mm, costal transverse section
prominent, semicircular; distal laminal cells 2-stratose, medial laminal
cells quadrate to rounded, straight, thin-walled; basal juxtacostal laminal
cells short-rectangular, straight, thin-walled; basal marginal laminal cells
short-rectangular, straight, with thick transverse and thin lateral walls,
not hyaline. Perichaetial leaves
not enlarged. Sexuality dioicous. Seta straight, 0.9--1.1 mm. Capsule commonly present, emergent,
chestnut brown, ovate with distinct constriction below rim, exothecial cells
rectangular, thick-walled, stomates absent, annulus of 1 row of quadrate,
thick-walled cells, operculum conic, peristome absent. Calyptra cucullate. Acidic sedimentary rocks; of conservation concern;
1900--2500 m; Grimmia nevadensis is a rare but locally abundant endemic, known only
from the 23. Grimmia serrana J. Muñoz, J. Shevock & D. R. Toren, J. Bryol.
24: 143. 2002 Plants in flat patches, olive green. Stems to 3 cm. Stem leaves
ovate-lanceolate from an ovate base, 2.5--3.5 × 0.6--0.7 mm, both margins
plane, costa-like intermarginal bands 2--4(--5)-stratose, awn to 1 mm, not
decurrent, narrowly attached, acuminate, costa narrow proximally, distal
laminal cells 1--3-stratose, oblate to rectangular, thick-walled; medial
laminal cells quadrate, slightly sinuose, thick-walled; basal juxtacostal
laminal cells short- to long-rectangular, sinuose, thick lateral-walled,
dense; basal marginal laminal cells quadrate to short-rectangular, sinuose,
thick lateral-walled, not hyaline. Perichaetial
leaves enlarged. Seta straight
to slightly curved, to 3.5 mm. Capsule
occasionally present, exserted, pale yellow, ovoid, exothecial cells oblong,
thin-walled, stomates absent, annulus of 1--2 rows of quadrate, thick-walled
cells, operculum short straight, peristome fully developed, not perforate,
not split. Calyptra cucullate. Humid to dry
areas, exposed granite, metamorphic rock, metavolcanic rocks and basalt,
montane woodlands; 670--1400 m; Grimmia serrana is endemic to western North America, having been
collected only in 24. Grimmia laevigata (Bridel)
Bridel, Bryol. Univ. 1(1): 183. 1826 Campylopus laevigatus Bridel, Muscol. Recent. Suppl. 4: 76. 1819 [1818]; Grimmia glauca Cardot; G. leucophaea Greville; G. sarcocalyx Kindberg Plants in hoary, dense tufts, dark green to dark brown. Stems 0.5--2 cm. Stem leaves oblong-ovate to oblong-lanceolate, 1.5--3 × 0.4--0.6
mm, both margins plane, intermarginal bands absent, awn 0.3--2 mm, decurrent,
broadly attached, acute, costa broad proximally; distal laminal cells
2-stratose, quadrate, thick-walled; medial laminal cells rounded-quadrate, straight,
thick-walled; basal juxtacostal laminal cells elongate, straight, thick
lateral walls, green; basal marginal laminal cells oblate to quadrate,
straight, thick transverse and thin lateral walls, green, not hyaline. Perichaetial leaves not enlarged. Seta straight, 1.5--3 mm. Capsule occasionally present,
exserted, brown, oblong-ovoid to cylindric, exothecial cells quadrate,
thick-walled, stomates present, annulus of 2--3 rows of rectangular,
thick-walled cells, operculum short rostrate, peristome irregularly perforate
distally, irregularly split. Calyptra
mitrate. Humid to dry,
exposed, acidic, sandstone and granite and basalt, open plains to montane,
rarely alpine; 240--2800 m; B.C.; Ala., Ariz., Ark., Calif., Colo., D.C.,
Fla., Ga., Idaho, Ill., Ind., Iowa, Kans., Ky., Md., Mass., Minn., Mo.,
Mont., Nebr., Nev., N.J., N.Mex., N.Y., N.C., Okla., Oreg., Pa., S.C.,
S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., Wyo.; Mexico; South America;
Eurasia; Africa; Indian Ocean Islands; Australia. Grimmia
laevigata is
widespread and relatively common on the southern Great Plains, into the
Ozarks and along the Appalachians from northeastern xxxd. Grimmia subg. Rhabdogrimmia Limpricht. Laubm. I: 759.
1889 Plants
1--8 cm. Stem central strand present or absent. Stem leaves
lanceolate to ovate-lanceolate, keeled, costa projecting on dorsal side, leaf
margin recurved on one or two sides; laminal cells usually 1-stratose distally
(2--4 stratose in G. elatior and G. funalis) except at margin; basal marginal cells not
hyaline (except G. funalis and G. torquata). Gemmae present or absent. Seta arcuate to cygneous when
moist, centrally attached to capsule. Capsule cernuous or pendulous,
striate to sulcate when dry and empty (smooth in G. attenuata), stomates
present at base of capsule (absent in G.
anomala); calyptra cucullate to mitrate. Species
32 (17 in flora): North America; Mexico; South America; Eurasia; Africa;
Pacific Islands; Australia. Members
of this subgenus are recognized by their lanceolate, thin, sharply keeled
leaves with recurved margins, typically 1-stratose lamina with 1- or
2-stratose margins, and long, arcuate seta with striate to sulcate capsules.
Most members of the subgenus occur on dry acidic rock. 25.
Grimmia lesherae H. C. Greven, Grimmias of the World. 130--131. 2003 Plants in
loose tufts, green. Stems 0.5--1.5 cm, dichotomously branched, central
strand absent. Stem leaves loosely appressed and straight when dry,
erectopatent when moist, broadly oblong-lanceolate, tapering to acute apex,
1.5--2 × 0.4--0.6 mm, larger towards stem tips, sharply keeled, margins
recurved on both sides, awns absent, occasionally short hyaline points at
leaf tips present, costa weak proximally, broad distally, projecting on
dorsal side; distal laminal cells
1-stratose, margins 2-stratose; medial
laminal cells short-rectangular with nodulose to sinuose walls, weakly
papillose; basal juxtacostal laminal
cells rectangular to linear, straight, thin-walled; basal marginal laminal cells rectangular,
thin-walled. Gemmae absent. Sexual
condition dioicous. Seta straight, 2 mm. Capsule
occasionally present, exserted, yellowish brown, oblate, smooth, exothecial
cells thin-walled, annulus present, operculum rostrate, peristome teeth
orange, deeply split and perforated, papillose. Calyptra cucullate. Damp
acidic rock; of conservation concern; 2000--3000 m; Grimmia
lesherae is apparently endemic to the high mountains along
the 26. Grimmia anomala
Schimper, Syn. Musc. Europ. ed. 2: 270. 1876 Grimmia
hartmanii ssp. anomala (Hampe) Loeske; G. pachyneurula
J.
K. A. Müller & Kindberg; G. philibertiana Britton Plants
in tufts, yellowish
green, blackish below. Stems 1.5--3.5 cm, small central strand
present. Stem leaves irregularly imbricate when dry, erect when moist,
oblong-lanceolate, gradually narrowed into a blunt chlorophyllose point,
1.5--2.5 × 0.4--0.8 mm, keeled, margins recurved on one or both sides, awns
absent to very short, costa projecting on dorsal side; distal laminal cells 1-stratose, in
places 2-stratose, margins 1-stratose; medial laminal cells rounded-quadrate,
walls slightly sinuose, thin- or thick-walled; basal juxtacostal laminal cells quadrate to short-rectangular, straight,
thin-walled; basal marginal laminal
cells quadrate to
short-rectangular with thickened transverse walls. Gemmae in clusters, globular, yellowish green to orange,
multicellular, on hyaline, deformed leaf apices. Sexual condition
dioicous. Seta straight to slightly curved when moist, 3--5 mm. Capsule
extremely rare, exserted, brownish, oblong-ovoid, smooth,
exothecial cells isodiametric, thick-walled, annulus present, operculum with
long straight beak, peristome teeth orange, fully developed, smooth below,
perforated and papillose distally. Calyptra
mitrate. Exposed,
damp acidic rock in boreal and alpine meadows and slopes; 200--3000 m;
Greenland; Alta., B.C., Nfld., N.S., Ont.; Alaska, Calif., Colo., Idaho,
Mont., Mich., Oreg., Utah, Wash., Wyo.; Eurasia. Because
of the always abundantly present gemmae on leaf apices, Grimmia anomala has often been confused with Grimmia hartmanii. Indeed, Grimmia anomala has frequently
been treated as a variety of G. hartmanii (G. N. Jones 1933, E. Lawton
1971). However, as pointed out by O. Vitikainen (1969), the ecological and
morphological features of this latter species are so constant and distinct,
that there is no doubt that it can be treated as a separate species. Special
features of G. anomala are stems with central strands, and
longitudinal ridges on cell walls in the distal part of the leaf that
resemble papillae in transverse section. It tends to prefer upland habitats.
In contrast, Grimmia hartmanii is principally a lowland species,
frequently growing on boulders in forests. It lacks a central strand and
longitudinal ridges are absent. Its spreading secund distal leaves taper to
long sharply keeled apices. 27. Grimmia attenuata (J.
K. A. Müller & Kindberg) Kindberg, Eur. N. Am. Bryin. 2: 228.
1897 Racomitrium alternuatum J. K. A.
Müller & Kindberg, Cat. Canad. Pl., Musci: 73. 1892 Plants robust,
brownish to reddish in distal part, black and frequently defoliated below. Stems 5--10 cm, central
strand absent. Stem leaves straight appressed when dry, erectopatent
when moist, lanceolate, 2.7--3.5 × 0.5--0.75 mm, sharply keeled distally,
margins narrowly reflexed on both sides, awns terete, firm and short,
denticulate, costa 70-120 µm wide at base, channeled distally, circular,
projecting on dorsal side; distal
laminal cells 1-stratose, apex and margins 2-stratose; medial laminal cells short- to long-
rectangular with extremely thick and sinuose lateral walls; basal juxtacostal laminal cells elongate with extremely thick,
slightly nodulose walls; basal marginal
laminal cells in a few
rows short-rectangular, thin-walled. Gemmae
absent. Sexual condition dioicous. Seta cygneous,
3--4 mm. Capsule sporadically present, exserted, yellowish brown,
ovoid, smooth, exothecial cells thin-walled with incrassate corners, annulus
present, operculum rostrate, peristome teeth orange, 40--60 µm wide at base,
split and perforated, smooth proximally, papillose distally. Calyptra not seen. Loose
tufts on dry boulders; of conservation concern; 0--1000 m; B.C.; This
northwestern North American endemic resembles forms of the extremely variable
Racomitrium heterostichum, and because G. N. Jones (1933) mentioned
five varieties of this latter species, it is not surprising that in some
herbaria, e.g. NY, all specimens of G. attenuata have been filed as
varieties of R. heterostichum. N. C. Kindberg (1897) realized that his
species was not a Racomitrium, but a Grimmia, and he wrote:
"Habit of G. elatior." In spite of Kindberg's correct
transfer to Grimmia, T. C. Frye (1918) synonymized it with R.
macounii. Adding to the confusion, H. Möller (1929) synonymized it with Grimmia
elatior, and J. Muñoz (1999b) published it as G. arcuatifolia
Kindberg, which is a synonym of G. lisae De Notaris. Grimmia
elatior is characterized by ellipsoid striate capsules, leaf margin
broadly recurved on just one side, and
an opaque 2-stratose distal lamina with rounded usually mammillose to
papillose cells, the mid-leaf cells quadrate to short-rectangular with
slightly sinuose, thick walls. 28. Grimmia brittoniae Williams,
Bull. Torr. Bot. Cl. 27: 316. 1900 Plants in
extremely hoary, compact glaucous cushions, brown inside. Stems 2--3
cm, central strand absent. Stem leaves loosely appressed to slightly
contorted when dry, erectopatent when moist, lanceolate, 0.5--1 × 0.3--0.4
mm, keeled, margins narrowly recurved on both sides, awns 2--4 mm, smooth,
flattened below, decurrent, costa weak, projecting on dorsal side; distal laminal cells 1-stratose,
margins 1-stratose; medial laminal
cells short-rectangular, slightly sinuose, thick-walled; basal juxacostal laminal cells rectangular,
straight to slightly sinuose, thick-walled; basal marginal laminal
cells quadrate with thickened
transverse walls. Gemmae absent. Sexual
condition dioicous. Seta flexuose, 2 mm. Capsule
occasionally present, exserted, brownish, ovoid, wide-mouthed, exothecial
cells thin-walled, annulus present, operculum conical, peristome teeth
yellow, split and perforated distally, slightly papillose. Vertical
faces of shaded, calcareous cliffs; of conservation concern; 500--700 m; Grimmia brittoniae is an endemic of western 29. Grimmia elatior Bals & De
Notaris, Mem. R. Acc. Sc. Torino 40: 340. 1838 Grimmia cognata Cardot &
Thérot; G. grandis Kindberg; G. papillosa (Warnstorf) Kindberg
Plants in
robust, dark green to blackish green, loose, hoary, readily disintegrating
tufts. Stems 1--5 cm, central
strand absent. Stem leaves loosely appressed to slightly twisted when
dry, erectopatent when moist, lanceolate to ovate-lanceolate, tapering to
acute apex, 2--3.0 × 0.5--0.7 mm, keeled, margin broadly recurved on one
side, awns short to long and weakly denticulate, costa weak at base,
channeled distally, projecting on dorsal side; distal laminal cells 2-stratose with thick, prominent
multistratose bands, margins multistratose and thick, areolation very opaque
with rounded thick-walled cells, occasionally papillose; medial laminal cells quadrate to
short-rectangular, sinuose, thick-walled; basal juxtacostal laminal cells short- to long-rectangular,
sinuose-nodulose, thick-walled; basal marginal
laminal cells quadrate to short-rectangular, thin- to thick-walled. Gemmae absent. Sexual condition
dioicous. Seta arcuate, 2--3
mm. Capsule occasionally present, emergent to exserted, brown, oblate,
striate, exothecial cells thin-walled, annulus present, operculum rostrate,
peristome teeth purple, deeply split, papillose. Calyptra mitrate. Exposed,
dry acidic rock and occasionally basic limestone; 500--4500 m; Greenland;
Alta., B.C., Yukon; Colo., Mont., N.J., Oreg., S.Dak., Wyo.; Eurasia; Africa. The rather robust Grimmia elatior was described by
Kindberg as Grimmia grandis. The
species is fairly common in the Canadian Rockies and in the western 30. Grimmia funalis (Schwägrichen)
Bruch and Schimper, Bryol. Eur. 3: 119. 1845 Trichostomum funale Schwägrichen, Sp. Musc. Frond. Suppl. 1(1): 150, tab 37. 1811; G. calvescens Kindberg; G. imberbis Kindberg; G. ryanii Limpricht; G. spiralis Hooker
Plants growing
in dense, usually hemispherical cushions, breaking up readily, falling apart
into clusters and string-like (spirally twisted leaves) single shoots,
greyish green. Stems 2--5 cm, central strand present. Stem leaves
usually spirally arranged when dry, patent when moist, lanceolate, 0.5--1.5 ×
0.2--0.5 mm, appressed, of ± uniform length throughout stem, keeled, margins
plane or recurved on one side, in female plants awns long and denticulate, in
male plants very short to absent, costa weak below, projecting on dorsal
side; distal laminal cells 2-stratose; medial laminal cells
short-rectangular, extremely sinuose, thick-walled; basal juxtacostal laminal cells elongate,
± sinuose, thick-walled; basal marginal
proximal laminal cells short-rectangular, hyaline. Gemmae absent. Sexual condition
dioicous. Seta arcuate, 1.5--2 mm. Capsule occasionally
present, exserted, yellowish-green, oblate, weakly striate, concealed in
awns, exothecial cells thin-walled, annulus present, operculum conical to
rostellate, peristome teeth orange, split distally, papillose. Calyptra mitrate. Damp acidic
rock; 500--2000 m; Nunavut, Ont.; La., Mich.; Eurasia. Grimmia
funalis is frequently misidentified because it is extremely
variable in height, color and length of the awns. In optimal conditions, it
forms extremely dense, greyish green cushions that break up easily, falling
apart into clusters and straight single plants. A distinct feature is the
string-like appearance of the dried up shoots, which results from the spiral
twisting of the leaves around the stem. However, H. C. Greven has also seen
populations without such spirally twisted leaves. Under the microscope, the
mid-leaf areolation with yellowish, short-rectangular, very thick and sinuose
cell walls is characteristic. Formerly recognized taxa Grimmia calvescens, G. imberbis and G. ryanii are
actually male plants of G. funalis. They grow in separate cushions
with muticous to short-awned leaves, deviating greatly from the much taller,
long-awned female plants. These male plants might be confused with G.
elongata, which frequently grows in the same habitat, or with G.
caespiticia. However, in G. elongata the basal cells are pellucid,
straight and thin-walled or only slightly incrassate. In G. caespiticia
the basal cells are shorter tending to be quadrate to short-rectangular, and
the leaf apex is cucullate. 31. Grimmia hartmanii Schimper, Syn. Musc. Eur.:
214. 1860 Plants in ascending, dichotomously
branched patches, green. Stems 2--8 cm, central strand absent. Stem
leaves slightly contorted with often secund apices when dry, distal
leaves usually falcate-secund when moist, broadly lanceolate, tapering to an
acuminate apex, 2.5--4.5 × 0.4--0.6 mm, keeled, margins usually recurved on
one side, awns very short, denticulate, costa firm, projecting on dorsal
side; distal laminal cells
1-stratose, margins 2-stratose; medial
laminal cells quadrate to short-rectangular, slightly sinuose,
thick-walled; basal juxtacostal
laminal cells short- to long-rectangular, slightly sinuose,
thick-walled; basal marginal laminal
cells quadrate, thickened transverse walls. Gemmae in clusters,
mulberry- or raspberry-shaped, brown, borne on leaf apices, usually present. Sexual
condition dioicous. [Seta straight to flexuose, 3--4 mm. Capsule
extremely rare, exserted, yellowish-green, clavate-oblate, smooth, exothecial
cells thick-walled, annulus present, operculum rostrate, peristome teeth
orange, deeply split, perforated distally, papillose. Calyptra mitrate.] Shaded boulders, especially
granite, in woodlands; 100--1500 m; P.E.I., Que.; Grimmia hartmanii is
principally a lowland species, occurring in the mountains up to 1500 m. With
the separation of G. anomala from this taxon, G. hartmanii is
much less common in 32.
Grimmia incurva Schwägrichen, Spec. Musc. Suppl. 1(1): 90. 1811 Grimmia
curvifolia Lindberg; G.
torngakiana Brassard & Hedderson Plants in
usually rounded cushions, green to blackish. Stems 1--2 cm, central strand present. Stem leaves
incurved and moderately contorted when dry, spreading when moist, oblong- to
linear-lanceolate, tapering to a slender and acuminate, often hyaline apex,
2.5--4.5 × 0.3--0.5 mm. keeled, margins plane or recurved below on one or
both sides, awns short, often only a hyaline point, occasionally long and
denticulate, costa projecting on dorsal side; distal laminal cells 1-stratose, margins and apex 2-stratose; medial laminal cells rectangular,
nodulose, thick-walled; basal juxtacostal
laminal cells long-rectangular, slightly sinuose, thick-walled; basal marginal laminal cells short-
to long-rectangular, thin-walled. Gemmae
absent. Sexual condition dioicous. Seta arcuate, 1.5--2 mm. Capsule
occasionally present, emergent to exserted, yellowish, oblate, smooth or
somewhat wrinkled when dry, exothecial cells thin-walled, annulus present,
operculum conic to rostrate, peristome teeth orange, divided distally,
papillose. Calyptra mitrate. Shaded
damp, acidic rock; 500--2500 m; Greenland; Alta., B.C., Nwfd. (Labr.), N.B.,
Yukon; Calif., Colo., Maine, N.H., N.Y, Oreg., S.Dak., Wash.; Eurasia. The peculiar east-west disjunct distribution of Grimmia
incurva in 33. Grimmia leibergii Paris, Ind.
Bryol. 528.
1896 Grimmia jacutica Plants robust,
ascending from a decumbent base, dark olive green. Stems 5--12 cm,
repeatedly dichotomous, central strand absent. Stem leaves loosely
appressed when dry, patent to spreading when moist, often becoming secund
distally on the stem, lanceolate to ovate-lanceolate, 3--4 × 0.6--0.9 mm,
keeled, margins recurved on both sides, awns denticulate, flattened
proximally, sometimes decurrent, costa yellowish to pale orange proximally,
wider (± 100 µm) near the base, channeled distally, semicircular on dorsal
side; distal laminal cells
1-stratose, margins partly 2-stratose; medial laminal cells quadrate to short-rectangular, strongly
sinuose, thin to thick oblique transverse walls and extremely thick lateral
walls; basal juxtacostal laminal
cells elongate to linear, weakly orange at insertion, nodulose,
thick-walled; basal marginal laminal cells in a few rows
quadrate to short-rectangular, thick-walled. Gemmae absent. Sexual condition dioicous. Seta arcuate, 3--4 mm. Capsule
occasionally present, exserted, yellowish-green, oblong-ovate, striate,
exothecial cells rather thick-walled, annulus present, operculum rostrate,
peristome teeth orange to reddish, perforated, irregularly cleft at apex,
nearly smooth basally, papillose. Calyptra
mitrate. Dry
acidic boulders; 400--500 m; B.C.; Grimmia leibergii, formerly thought
to be endemic to western 34. Grimmia lisae De Notaris,
Musc. Ital. Spic. 15. 1837 Grimmia ancistrodes Durieu &
Montagne; G. arcuatifolia Kindberg; G. californica Sullivant; G.
canadensis Kindberg; G. flettii (Holzinger) Cardot Plants in
dense to loose tufts, olive green, brownish to blackish below. Stems 1--4
cm, central strand present. Stem leaves erect and appressed when dry,
recurved to squarrose when moist, broadly lanceolate, tapering to an acute
apex, 1.5--2.5 × 0.4--0.6 mm, keeled, margins recurved on one or both sides,
awns absent to rather long, stout and denticulate, costa reniform, projecting
on dorsal side, median layer of stereids present; distal laminal cells 1-stratose with 2-stratose ridges, margins
2-stratose; medial laminal cells
rounded-quadrate to oblate, straight, thick-walled; basal juxtacostal laminal cells
short-rectangular to occasionally elongate, straight to slightly sinuose,
thin- to thick-walled; basal marginal
laminal cells quadrate to short-rectangular, thickened transverse
walls. Gemmae in clusters,
multicellular, occasionally present in leaf axils. Sexual condition
dioicous. Seta arcuate, 3--4.5 mm. Capsule occasionally present,
exserted, ovoid, brown, shiny, weakly striate, exothecial cells thin-walled,
annulus present, operculum rostrate, peristome teeth orange, fully developed
to irregularly cleft at apex, papillose. Calyptra
mitrate. Dry
acidic to basic rock; 60--1000 m; B.C.; Calif., Oreg., Wash.; Mexico; n
Africa; Eurasia; Pacific Islands (Hawaii). Grimmia
lisae is a thermophilous species with a preference for subtropical coastal
areas. In North America, it occurs along the west coast, from Vancouver
Island south to 35. Grimmia moxleyi Holzinger,
Musci Bor. Am. Eur. 24. 1926 Plants in
frequently extended mats, blackish green. Stems 1--1.5 mm high, small central strand present. Stem
leaves erect with slightly incurved tips when dry, erect-spreading when
moist, oblong, broadly rounded and muticous at apex, 1.5--2 × 0.4--0.6 mm,
keeled, margins plane to recurved, awns absent, only present in perichaetial
leaves, costa weak below, projecting at dorsal side, perichaetial leaves
longer and with awns. Distal laminal
cells 2-stratose; medial
laminal cells rounded-quadrate, ± sinuose, thin- or thick-walled;
basal juxacostal laminal cells
short- to long-rectangular, straight, thin-walled; basal marginal laminal cells short- to long-rectangular,
thin-walled. Gemmae absent. Sexual
condition autoicous. Seta flexuose to curved,
1--1.5 mm. Capsule usually present, exserted, chestnut
brown, oblong-ovoid, wrinkled-plicate when dry, exothecial cells thin-
to thick-walled, annulus present, operculum conical, peristome teeth
yellowish, split and perforated, papillose. Calyptra cucullate. Dry acidic rock; of
conservation concern; 500--1500 m; Grimmia
moxleyi is endemic to the southwestern 36. Grimmia muehlenbeckii Schimper, Syn. Musc. Eur.: 212. 1860 Grimmia hermannii H. A. Crum; G.
trichophylla var. tenuis (Wahlenberg) Wijk & Margadant Plants in blackish green tufts. Stems
1--2.5 cm, central strand present. Stem leaves loosely appressed,
twisted when dry, erectopatent when moist, ovate-lanceolate, tapering to
acute apex, 2--3 × 0.6--0.8 mm, keeled, margins recurved in mid-leaf on both
sides, awns short, denticulate, in perichaetial leaves stout and often
decurrent, costa channeled above, projecting at dorsal side, angled to
bluntly winged; distal laminal cells
1-stratose with 2-stratose ridges, margins 2-stratose; medial laminal cells quadrate to
short-rectangular, sinuose, thick-walled; basal juxtacostal laminal cells short- to long-rectangular, yellowish,
nodulose, thick-walled; basal
marginal laminal cells short-rectangular with thickened transverse
walls. Gemmae rare, in clusters,
short-stalked, in distal leaf axils. Sexual condition dioicous. Seta
arcuate, 2--3 mm. Capsule occasionally present, exserted, globose,
shiny, brown, smooth to slightly striate, exothecial cells thin-walled,
annulus present, operculum rostrate, peristome teeth purple, fully developed
or slightly split distally, papillose. Calyptra
mitrate. Shaded acidic rock, often
along lakes; 200--2000 m; Nfld., Ont.; Grimmia muehlenbeckii is closely related to G.
trichophylla, and in the past it was frequently conceived as being a
subspecies or variety of that taxon. H. Deguchi (1978) treated it as a
distinct species, followed by A. J. E. Smith (1992). H. C. Greven agrees with
these treatments. He has seen many specimens that are remarkably uniform and
easy to distinguish from G. trichophylla by their small, globose,
shiny, dark brown capsules with purple, entire peristome teeth. In contrast,
the capsules in G. trichophylla are oblong-ovoid, larger and longer
than in G. muehlenbeckii, dull,
yellowish-brown, and the peristome teeth are orange and cleft. The
gametophyte differs from G. trichophylla by its blackish green tufts,
angled costa with blunt wings protruding on the dorsal side, and
ovate-lanceolate leaves with stout denticulate awns that are often decurrent. 37. Grimmia orbicularis Wilson,
Engl. Bot. Suppl. 4: 2888. 1844 Plants in
hoary, usually hemispherical cushions, greyish green. Stem 2--5 cm,
central strand present. Stem leaves appressed and twisted when dry,
erect when moist, broadly lanceolate, abruptly contracted into hair-point,
2--2.5 × 0.4--0.6 mm, keeled, margins recurved in the middle of the
leaf on one or both sides, awns short to long, smooth to denticulate, costa
weak below, projecting on dorsal side; distal laminal cells 1-stratose; medial laminal cells subquadrate, sinuose, thick-walled; basal juxtacostal laminal cells elongate,
nodulose, thick-walled; basal marginal
laminal cells short- to long-rectangular with thickened transverse
walls. Gemmae absent. Sexual condition autoicous. Seta arcuate, 2--3 mm. Capsule usually present,
exserted, bent down into the cushions by the arcuate setae, yellowish brown
to chestnut brown, globose, smooth, when dry and empty wide-mouthed, wrinkled
and ribbed, exothecial cells thin-walled,
annulus present, operculum mammillate, peristome teeth orange, broad,
cribrose and irregularly cleft at apex. Calyptra
cucullate. Dry
basic rocky substrates such as limestone, basalt, and mortar; 200--2000 m; Grimmia
orbicularis is a thermophilous species with a preference for
sunny, basic substrates. In 38. Grimmia pulvinata (Hedwig)
Smith, Engl. Bot. 24: 1728. 1807 Fissidens pulvinatus Hedwig, Sp. Musc.
Frond.: 158. 1801; Grimmia decipiens var. hendersonii (Renald &
Cardot) Sayre; G. indianensis (Sayre) Crum; G. pulvinata var.
africana J. D. Hooker f. & Wilson; G. subcurvula Kindberg; G.
trichophylla var. indianensis
Sayre Plants in
hemispherical hoary cushions, greyish green. Stems 1--3 cm, central
strand present. Stem leaves flexuose when dry, erect when moist,
lanceolate, 1--1.7 × 0.3--0.6 mm, keeled, margin recurved on both sides
nearly from base to apex, abruptly contracted into short to long, smooth to
denticulate awn, costa weak proximally, projecting on dorsal side; distal laminal cells 1-stratose,
margins 2-stratose; medial laminal
cells rounded-quadrate, slightly sinuose, thin-walled; basal juxtacostal laminal cells
short-rectangular, thin-walled; basal marginal
laminal cells quadrate, thin-walled. Gemmae absent. Sexual
condition autoicous. Seta arcuate, 3--4 mm. Capsule
usually present, exserted, ellipsoid to oblate, brownish and ribbed when
empty and dry, exothecial cells thin-walled, annulus present, operculum
rostrate, peristome teeth reddish, fully developed, papillose. Calyptra mitrate. Various
substrates, from acidic to basic rock, old mortar, tree trunks; 0--3000 m;
Alta., B.C., Ont.; Ariz, Calif., Colo., Idaho, Ind., Iowa, Kan., Md, Mich.,
Miss., Mo., Mont., Nev., N.Mex., Okla., Ore., S.Dak., Tex., Utah, Wash.,
Wyo.; Mexico; South America (Argentina, Chile, Uruguay); Eurasia; Africa;
Pacific Islands (New Zealand); Australia. Grimmia
pulvinata is the most common Grimmia species. It has a
nearly cosmopolitan distribution, and is a pioneer on various substrates,
even on the trunks of trees. However, in the eastern part of 39. Grimmia ramondii (Lamarck & De Candolle)
Margadant, Lindb. 1: 128. 1972 Pterigynandrum ramondii Lamarck & De Candolle, Fl. Franc. ed. 3 [original issue] 2: 462.
1805; Dryptodon patens (Hedwig)
Bridel; Grimmia curvata (Bridel) De Sloover, G. serrata
Kindberg; Racomitrium patens
(Hedwig) Huebener Plants in
robust, loose, arched-ascending, readily disintegrating patches, brownish
green distally, blackish below. Stems 5--10 cm, central strand
absent. Stem leaves loosely appressed when dry, patent when moist,
broadly ovate-lanceolate, tapering to an acute, slightly toothed apex, 2--3 ×
0.4--0.6 mm, keeled, margins recurved on both sides, awns absent, costa
stout, with two dorsal lamellae (or wings) forming parallel ridges along its
length; distal laminal cells
1-stratose, margins 2-stratose; medial
laminal cells quadrate to rectangular, sinuous, thick-walled; basal juxtacostal laminal cells yellow,
linear, sinuose, thick-walled; basal marginal
laminal cells quadrate to short-rectangular, slightly thick-walled. Gemmae absent. Sexual condition
dioicous. Seta cygneous to
arcuate at maturity, flexuose when old, 3--5 mm. Capsule
occasionally present, exserted, oblate, yellowish green to yellowish brown,
plicate when empty, exothecial cells thin-walled, annulus present, operculum
rostrate, peristome teeth purple, divided nearly to base into two partly
adhering segments, papillose. Calyptra
mitrate. Dry and
damp acidic rock; 0--2000 m; Greenland; Alta., B.C. Nfld., N.W.T.; Alaska,
Calif., Idaho, Mich., Mont., Oreg., Wash.; Europe; Asia (China, Japan); n
Africa. Grimmia
ramondii occurs near sea level in Arctic areas and along the
West Coast. In the latter region, however, it is more common in mid-elevation
montane sites and may also be found above the tree line. Grimmia ramondii
differs from most Grimmia species by its lack of awns, its winged
costa and tall, loose growth form. At first sight, G. ramondii has the
habit of a Racomitrium. However, the sporophyte has an arcuate rather
than a straight seta and the capsules are striate rather than smooth. Because
of its habit and the absence of a central strand, G. K. Limpricht (1890)
placed it, together with G. hartmanii and G. atrata in Dryptodon,
a genus intermediate between Grimmia and Racomitrium. There are
many other Grimmia species, however, without a central strand, and G.
hartmanii and G. atrata have already been removed from Dryptodon,
so there is no reason to maintain this monotypic genus. Grimmia ramondii
may be confused with G. hartmanii, and G. elatior. However, the lack of awns, the Racomitrium-like
areolation, and the distinct costal wings distinguishes it easily from these
two species. 40. Grimmia torquata Drummond,
Musci Scotici, Vol. 2: 28. 1825 Grimmia
pellucida Kindberg; G. prolifera J. K. A. Müller &
Kindberg; G. pseudotorquata Kindberg; G. tortifolia Kindberg Plants in
soft, readily disintegrating cushions, yellow-green to brownish, occasionally
light green, blackish to red-brown inside. Stems 1--4 cm, small
central strand present. Stem leaves contorted when dry, patent when
moist, lanceolate, 1.5--2 × 0.3--0.5 mm, keeled, margins slightly recurved
below, plane above, awns very short and smooth, occasionally absent, costa
weak below, projecting on dorsal side; distal laminal cells 1-stratose; medial laminal cells rectangular, extremely sinuose,
thick-walled; basal juxtacostal
laminal cells linear, extremely sinuose, thick-walled; basal marginal laminal cells long rectangular,
thin-walled, hyaline. Gemmae
brown, multicellular, present on the dorsal side of distal leaves. Sexual
condition dioicous. Seta slightly curved, straight when dry,
3--5 mm. Capsule sporadically present, exserted, ovoid, brown, smooth,
striate when dry and empty, exothecial cells thin-walled, annulus present,
operculum rostrate, peristome teeth yellowish, short, split in distal part,
weakly papillose. Calyptra
mitrate. Damp,
frequently vertical faces of acidic rock; 200--4000 m; Greenland; Alta.,
B.C., Labrador, Nfld., N.W.T., Ont., Que., Yukon; Alaska, Ariz., Calif.,
Colo., Idaho, Mont., Nev., Ore., Wash., Wyo.; Mexico; n Africa; Eurasia;
Pacific Islands (Hawaii). Grimmia
torquata is a montane-alpine species with a preference for
shaded habitats. It grows in hemispherical cushions on steep, damp rock walls.
Preferred substrates are granite, gneiss, quartzite and schist. Its
distribution reaches from near sea level in the 41. Grimmia trichophylla Greville,
Fl. Edinensis: 235. 1824 Plants in dense to loose patches,
yellowish green to dark green. Stems 2--4 cm, central strand present. Stem
leaves loosely appressed, slightly twisted when dry, erectopatent when
moist, lanceolate to oblong-lanceolate, tapering to acute apex, 2--3.5 ×
0.3--0.4 mm, usually sharply keeled, margins recurved on one or both sides,
plane to erect distally, awns variable, short to long, smooth to denticulate,
not conspicuously flattened at base, costa firm, projecting on abaxial side; distal laminal cells 1-stratose,
occasionally with 2-stratose ridges; medial
laminal cells quadrate to short-rectangular, slightly sinuose,
thick-walled; basal juxtacostal
laminal cells long-rectangular (rarely short rectangular), ± nodulose,
thick-walled; basal marginal laminal
cells short- to long-rectangular, with thickened transverse walls. Gemmae clusters occasionally present
in distal leaf axils. Sexual condition dioicous. Seta arcuate, 2--4 mm. Capsule
occasionally present, exserted, oblong-ovoid, yellowish green to stramineous,
striate when dry, exothecial cells thin-walled, annulus present, operculum
rostrate, peristome teeth yellowish, papillose, deeply split and perforated. Calyptra mitrate. Dry, acidic rock; 200--2000
m; B.C.; Ariz., Calif., Colo., Idaho, Maine, Mo., Mont., Nev., Okla., Oreg.,
S.Dak., Utah, Vt., Wash., Wyo; Mexico; Eurasia; Pacific Islands (Hawaii);
Australia. In xxxe. Grimmia subg. undetermined 42. Grimmia mollis Bruch & Schimper, Bryol. Eur. 3: 133. 1849 Hydrogrimmia mollis (Bruch & Schimper) Loeske; Grimmia
evansii E. Britton Plants in
loose
patches, green. Stems 1--5 cm,
central strand present. Stem leaves oblong to ovate-lanceolate, 2--3 ×
0.8--1 mm, margins plane, incurved above, muticous, concave, apex rounded to
cucullate, costa not projecting on dorsal side, ending well below apex;
juxtacostal and marginal distal
laminal cells 1-stratose; medial laminal cells quadrate, thin-walled;
basal juxtacostal laminal cells quadrate, thin-walled;
basal marginal laminal cells quadrate, thin-walled. Sexual condition dioicous. Seta
straight, 2--3 mm, yellowish. Capsule sporadically present,
exserted, yellowish, ovoid to oblong, smooth, exothecial cells thick-walled,
annulus absent, operculum rostellate, peristome teeth purple, perforated,
fully developed, papillose. Calyptra
cucullate. Wet acidic rocks in alpine
and boreal habitats; 1000--4115 m.; Greenland; Alta, B.C., Grimmia mollis, discovered by Schimper in
the Tirol region of 43. Grimmia
shastae Greven, Grimmias of the World. 208--209. 2003 Plants in
hoary tufts, brownish green. Stems
0.3--1 cm. Stem leaves ovate to broadly oblong-lanceolate, 1--1.5 ×
0.5--0.6 mm, larger towards stem tips, not keeled, frequently plicate in
extreme apex, margins plane, incurved above, long-awned, decurrent, costa
weak below, broadened and deeply channeled above; distal laminal cells 1-stratose with 2-stratose ridges, in apex
entirely 2-stratose; medial laminal cells
short-rectangular, sinuose, slightly thick-walled; basal juxtacostal laminal cells
rectangular, straight, with thin walls; basal marginal laminal cells rectangular, with thick transverse walls
and thin lateral walls, hyaline in 3--4 rows. Gemmae clusters of globular, brown, multicellular gemmae
abundantly present on the adaxial side of the distal lamina. Sexual
condition probably dioicous, gametangia not seen. Capsule unknown. Volcanic
rock; of conservation concern; 3500 m; Grimmia
shastae is a rare endemic known only from OTHER
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