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BFNA | Family List | BFNA Vol. 1 | Grimmiaceae

Grimmia Hedwig, Sp. Musc. Frond. 75. 1801.
[For Dr. J.F.K. Grimm, physician and botanist of Gotha, Germany (d. 1821)]

Authors: Roxanne I. Hastings & Dr. Henk C. Greven

  • Dryptodon Bridel
  • Hydrogrimmia (I. Hagen) Loeske

    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 94 (44 in the flora): North America, Mexico, Central America, South America, Eurasia, Africa, Pacific Islands, Australia, Antarctic.

    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, 20--24 and 44; 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 Southern Peru: 19--74. Tokyo. Greven, H. C. 1999. Synopsis of Grimmia in Mexico, including Grimmia mexicana, sp. nov. Bryologist 102: 426--436. Greven, H. C. 2002. Grimmia nevadense, a new species from California. Bryologist 105: 273--275. Greven, H. C. 2003. Grimmias of the World. Backhuys Publishers. Leiden. The Netherlands. Ireland, R. R. and Miller, N. G. 1982. Grimmia anodon (Musci; Grimmiaceae) in North America north of Mexico. Bryologist 85: 112--114. Muñoz, J. 1997. The correct name of Grimmia alpestris (Musci, Grimmiaceae). Bryologist 100: 517--519. Ochyra, R. 1993. Grimmia plagiopodia (Musci, Grimmiaceae) in the Southern Hemisphere. Fragm. Flor. Geobot. 38: 21--27. Sayre, G. 1952. Key to the species of Grimmia in North America. Bryologist 55: 251--259. Weber, W. A., R. C. Wittmann and R. Worthington. 2003. Grimmia bernoullii Mueller Hal. in the United States. Evansia 20: 104--106.

    OTHER REFERENCES

    Cao, T. and D. H. Vitt. 1986. A taxonomic revision and phylogenetic analysis of Grimmia and Schistidium (Bryopsida; Grimmiaceae) in China. J. Hattori Bot. Lab. 61: 123--247.
    Corley, M. F. V., A. C. Crundwell, A. C. Düll, R. Hill & A. J. E. Smith. 1981. Mosses of Europe and the Azores; an annotated list of species with synonyms from the recent literature. J. Bryology 11: 609--689.
    Crum, H. A. 1977. Grimmia pulvinata in Eastern North America. Bryologist 80: 152--153.
    Crum, H. A. 1994. Grimmiales. In: A. J. Sharp, HA. Crum, & P. M. Eckel (eds.). The Moss flora of Mexico. Part One. Sphagnales to Bryales. Mem. New. York Bot. Gard. 69: 386--415.
    Crum, H. A. and L. E. Anderson. 1981. Mosses of Eastern North America. 2 vols. Columbia University Press, N. Y.
    Deguchi, H. 1978. A revision of the genera Grimmia, Schistidium and Coscinodon (Musci) of Japan. J. Sci. Hiroshima Univ. Ser. B., Div. 2, Bot. 16: 121--256.
    Frye, T. C. 1918. The Rhacomitriums of western North America. Bryologist 21: 1--16.
    Greven, H. C. 1995. Grimmia Hedw. (Grimmiaceae, Musci) in Europe. Leiden. The Netherlands.
    Greven, H. C. 1997. Grimmia austro-funalis C. Müll., a species with a misleading name and a disjunct distribution. J. Bryology 19: 829--832.
    Greven, H. C. 1998. Grimmia Hedw. in Iceland, including Grimmia grisea Cardot, new to Europe. Lindbergia 23: 91--93.
    Greven, H. C. 1999. A synopsis of Grimmia in Mexico, including Grimmia mexicana, sp. nov. Bryologist 102: 426--436.
    Greven, H. C. & T. Spribille. 1999. Grimmia brittoniae, a rare moss endemic to northwestern Montana. Bryologist 102: 116--118.
    Grout, A. J. 1933. Moss Flora of Norh America. Newfane, Vermont. Vol. II. Newfane, Vermont.
    Hagen, I. 1909. Forarbejder til en norsk lovmosflora, IX--XII. K. Norske Vidensk. Selsk. Skrifter, Trondhjem.
    Hastings, R. I. 2002. Biogeography of Grimmia teretinervis (Bryopsida, Grimmiaceae) in North America. Bryologist 105: 262--266.
    Ireland, R. R. 1982. Grimmia teretinervis (Musci: Grimmiaceae) in North America. Bryologist 85: 332--334.
    Ireland, R. R., C. D. Bird, G. R. Brassard, W. B. Schofield, and D. H. Vitt. 1980. Checklist of the Mosses of Canada. Publ. in Botany No. 8. National Museums of Canada, Ottawa.
    Ireland, R. R., G. R., Brassard, W. B. Schofield, and D. H. Vitt. 1987. Checklist of the mosses of Canada II. Lindbergia 13: 1--62.
    Jones, G. N. 1933. Grimmia. In: A.J. Grout (ed.), Moss Flora of North America 2(1): 1--66. Newfane,Vermont.
    Keever, C., H. J. Oosting, and L. E. Anderson. 1951. Plant succession on exposed granite of Rocky Face Mountain, Alexander Co., North Carolina. Bull. Torrey Bot. Club 78: 401--421.
    Kindberg, N. C. 1897. Species of European and Northamerican Bryinae (Mosses) Part 2 Acrocarpous. Linköping.
    Lawton, E. 1971. Moss flora of the Pacific northwest. Hattori Botanical Laboratory. Nichinan, Japan.
    Leiberg, J. B. 1893. Two new species of mosses from Idaho. Bull. Torrey Bot. Club. 20: 113--115.
    Limpricht, G. K. 1890. Die Laubmoose, Deutschlands, Oesterreichs und der Schweiz. Vol. 1. Leipzig.
    Loeske, L. 1913. Die Laubmoose Europas, I. Grimmiaceae. M. Lande, Berlin.
    Maier, E. 2002. The genus Grimmia (Musci, Grimmiaceae) in the Himalaya. Candollea 57: 143--238.
    Möller, H. 1929. A revision of some new species and varieties of Rhacomitrium described by N. C. Kindberg from North America. Revue Bryologique 2(56): 82--86.
    Muñoz, J. 1998a. Materials toward a revision of Grimmia (Musci, Grimmiaceae): nomenclature and taxonomy of Grimmia longirostris. Ann. Miss. Bot. Gard. 85: 352--363.
    Muñoz, J. 1998b. A taxonomic revision of Grimmia subgenus Orthogrimmia (Musci, Grimmiaceae). Ann. Miss. Bot. Gard. 85: 367--403.
    Muñoz, J. 1999a. A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Ann. Miss. Bot. Gard. 86: 118--191.
    Muñoz, J. 1999b. Grimmia arcuatifolia and G. leibergii (Musci, Grimmiaceae), two neglected species from Northwestern North America. Ann. Jardín Botánico de Madrid 57: 7--13.
    Muñoz, J. 2000. New synonyms in Grimmia (Grimmiaceae). J. Bryology 22: 99--102.
    Muñoz , J. and Pando, F. 2000. A world synopsis of the genus Grimmia. Missouri Bot. Gard. Monog. 83. 133p.
    Muñoz, J., J. R. Shevock and D. Toren. 2002. Grimmia serrana (Bryopsida, Grimmiaceae), a new species from California, U.S.A. J. Bryol. 24: 143--146.
    Norris, D. H. and J. R. Shevock. 2004. Contributions toward a bryoflora of California: I. A specimen-based catalogue of mosses. Madroño 51: 1--131.
    Ochyra, R. 1993. Two new species of Grimmia from New Zealand. Frag. Flor. Geobot. Suppl. 2: 219--227.
    Oosting, H. J. and L. E. Anderson. 1937. The vegetation of a barefaced cliff in western North Carolina. Ecology 18: 280--292.
    Oosting, H. J. and L. E. Anderson. 1939. Plant succession on granite rocks in eastern North Carolina. Bot. Gaz. 100: 750--768.
    Sainsbury, G. O. K. 1945. New and critical species of New Zealand mosses. Trans. Royal Soc. New Zealand Vol. 75, Part 2: 169--186.
    Sayre, G. 1951. The identity of Grimmia ovalis and Grimmia commutata. Bryologist 54: 91--94.
    Sayre, G. 1955. Grimmia mariniana, a new species from California. Bryologist 58: 323--325.
    Smith, A. J. E. 1992. The taxonomic status of the British varieties of Grimmia trichophylla Grev. J. Bryology 17: 269--273.
    Vitikainen, O. 1969. On the taxonomy and distribution of Grimmia anomala and G. hartmanii Schimper. Ann. Bot. Fenn. 6: 236--242.

    1 Costa ending well below the apex; leaf tip rounded to cucullate, muticous; lamina uniformly 1-stratose; laminal cells all quadrate.   Grimmia mollis
    + 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 (1) Seta eccentrically attached to capsule base; capsule immersed, smooth, ventricose; stomates 3--4, large, at base of capsule.   Grimmia subg. Gasterogrimmia
    + 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)
           
    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)
    + Distal leaves concave; distal lamina 2-stratose with 2-stratose margins; annulus prominent; operculum rostrate.   (5)
           
    4 (3) Peristome present, annulus absent; distal lamina 1-stratose, margins 1- or 2-stratose.   Grimmia plagiopodia
    + Peristome absent, annulus present; distal lamina 1-stratose with 2-stratose patches, margins 2-stratose.   Grimmia anodon
           
    5 (3) Basal laminal cells thick-walled; gonioautoicous; peristome fully developed.   Grimmia americana
    + Basal laminal cells thin-walled; dioicous; peristome rudimentary.   Grimmia poecilostoma
           
    6 (2) Leaves 2--3-stratose distally or 1-stratose with distal margins widely 2-stratose (G. sessitana and some G. donniana and G. afroincurva); leaf margins incurved, plane or recurved.   (7)
    + Leaves merely 1-stratose distally; margins at most narrowly 2-stratose; one or both leaf margins recurved (plane in G. reflexidens) (mostly subg. Rhabdogrimmia).   (36)
           
    7 (6) Leaves concave; costa not prominent; margins plane or incurved; dioicous (subg. Litoneuron).   (8)
    + Leaves keeled; costa prominent; margins recurved, plane or incurved; autiocous or dioicous (mostly subg. Guembelia).   (14)
           
    8 (7) All leaves muticous, straight or falcate.   (9)
    + At least distal leaves awned, straight.   (10)
           
    9 (8) Leaves oblong-lanceolate to ligulate, straight, cucullate, obtuse-rounded; basal marginal laminal cells short-rectangular; widely distributed.   Grimmia unicolor
    + Leaves oblong-lanceolate, homomallous-falcate, subulate, uncinate; basal marginal laminal cells quadrate to short-rectangular; endemic to Pacific Coast region   Grimmia hamulosa
           
    10 (8) Leaves oblong-ovate to oblong-lanceolate; basal marginal laminal cells oblate to quadrate; costa broad at base; awn broadly attached and decurrent.   Grimmia laevigata
    + 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)
           
    11 (10) Costa-like multistratose leaf bands present in transverse section.   Grimmia serrana
    + Costa-like multistratose leaf bands absent.   (12)
           
    12 (11) Capsule emergent, gymnostomous, stomates absent, operculum mammilate; basal laminal cells uniformly short-rectangular, straight and thin-walled.   Grimmia nevadensis
    + Capsule exserted, peristome fully developed, stomates present, operculum rostrate; basal marginal and juxtacostal laminal cells typically contrasting in length or thickness.   (13)
           
    13 (12) 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.   Grimmia ovalis
    + 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.   Grimmia olneyi
           
    14 (7) Margins recurved on one or both sides.   (15)
    + Margins plane or incurved.   (27)
           
    15 (14) Leaves muticous, cucullate.   Grimmia atrata
    + Leaves awned, not cucullate.   (16)
           
    16 (15) Leaves narrowly lanceolate to linear-lanceoalate, contorted when dry; basal region entirely hyaline.   Grimmia afroincurva
    + Leaves lanceoate to oblong-lanceolate, straight or spirally curved around stem when dry; basal region not hyaline or only basal marginal cells hyaline.   (17)
           
    17 (16) Leaves spirally curved around stem when dry, flagelliform innovations present.   Grimmia funalis
    + Leaves straight when dry, flagelliform innovations absent.   (18)
           
    18 (17) Sporophytes present.   (19)
    + Sporophytes absent.   (23)
           
    19 (18) Capsule exserted.   (20)
    + Capsule immersed.   (22)
           
    20 (19) Seta arcuate; dioicous.   Grimmia elatior
    + Seta straight; autoicous.   (21)
           
    21 (20) Annulus prominent, 2 rows of rectangular cells; stomates in 2--3 rows; basal juxtacostal cells sinuose and thick walled   Grimmia longirostris
    + Annulus small, 1 row of quadrate cells; stomates in 1 row; basal juxtacostal cells straight and thin-walled.   Grimmia sessitana
           
    22 (19) Stem central strand present, epidermis thin; leaf margins 2-stratose, not thickened, one margin recurved (rarely both); leaves broadly ovate-lanceolate   Grimmia arizonae
    + 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   Grimmia pilifera
           
    23 (18) Basal juxtacostal cells straight, thin-walled; distal juxtacostal lamina 1-stratose, cells often bulging; plants small (less than 1 cm), blackish; moist, alpine habitats.   Grimmia sessitana
    + Basal juxtacostal cells sinuose, thick-walled; distal juxtacostal lamina at least 2-stratose, cells not bulging; plants robust (more than 1 cm), yellow-green to very dark olive green; dry, widely distributed.   (24)
           
    24 (23) Leaf margins 2-stratose, not thickened; stem central strand present, epidermis thin.   (25)
    + Leaf margins multistratose and thickened; stem central strand absent, epidermis thick.   (26)
           
    25 (24) Autoicous; costa transverse section reniform; leaves sheathing; widely distributed.   Grimmia longirostris
    + Dioicous; costa transverse section semicircular; leaves not sheathing; American Southwest   Grimmia arizonae
           
    26 (24) Leaves narrowly lanceolate from an ovate base, usually narrowly recurved on both margins; distal lamina lacking multistratose bands, never papillose   Grimmia pilifera
    + Leaves broadly lanceolate, broadly recurved on one margin; distal lamina with multistratose bands, occasionally papillose.   Grimmia mariniana
           
    27 (14) Costa transverse section circular distally; awn hyaline-tipped to short (0.3 mm), decurrent.   Grimmia teretinervis
    + osta transverse section semicircular distally; awn short to long, usually prominent, decurrent or not.   (28)
           
    28 (27) Seta arcuate; leaves spirally curved around stem when dry, flagelliform innovations present.   Grimmia funalis
    + Seta straight; leaves straight or contorted when dry but not curved around stem, flagelliform innovations absent.   (29)
           
    29 (28) Gemmae abundant on adaxial distal leaf surface/   Grimmia shastae
    + Gemmae absent on leaves.   (30)
           
    30 (29) Distal laminal cells not bulging.   (31)
    + Distal laminal cells bulging.   (36)
           
    31 (30) Margins plane throughout; basal marginal laminal cells short- to long-rectangular; stomates present; autoicous.   (32)
    + Margins incurved distally; basal marginal laminal cells quadrate to short-rectangular; stomates absent; dioicous.   (34)
           
    32 (31) Leaves straight, ovate-lanceolate to oblong-lanceolate; basal marginal cells hyaline or not.   (33)
    + Leaves contorted, narrowly lanceolate; entire basal region hyaline.   Grimmia afroincurva
           
    33 (32) Distal juxtaxcostal laminal cells (1--)2-stratose; basal marginal leaf cells long-rectangular, hyaline; annulus large, of 2 rows of rectangular cells, revoluble; calyptra mitrate   Grimmia donniana
    + 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   Grimmia sessitana
           
    34 (31) Capsule immersed to emergent, wide-mouthed, peristome rudimentary; endemic to California.   Grimmia mariniana
    + Capsule exserted, narrow-mouthed, peristome present, fully developed; widespread in western North America.   (35)
           
    35 (34) Basal juxtacostal laminal cells short- to long-rectangular, distinct from quadrate to short-rectangular basal marginal cells; medial laminal cells rounded, thick-walled.   Grimmia montana
    + Basal juxtacostal and marginal cells quadrate to short-rectangular, basal areolation relatively uniform; medial laminal cells quadrate to short-rectangular, thin-walled.   Grimmia alpestris

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