Common Names in English:
Plants acrocarpous or cladocarpous
, small to large, usually olivaceous
to blackish green, growing in rigid
, tufts, mats or patches. Stems erect
, or prostrate
, dichotomously to irregularly branched. Leaves erect and tightly appressed
when dry, erect-spreading to patent
when wet, lanceolate to ovate-lanceolate, less often ovate
, oblong-ovate, linear
, or lingulate
, to broadly concave
or sometimes longitudinally plicate
, rarely with adaxial
lamellae (Indusiella), margins
, or variously recurved or revolute
, mostly entire, 1- to multistratose, acuminate, acute to rounded-obtuse, typically with a hyaline
, sometimes muticous
, costa single, rarely spurred
distally (Codriophorus and Niphotrichum), usually strong
to excurrent, rarely subpercurrent, typically with one stereid
, distal lamina 1-2(-4) -stratose; basal cells
, rarely oblate
, straight, sinuose, or nodulose
, basal juxtacostal and marginal
regions usually differentiated, alar cells
undifferentiated or hyaline; mid leaf cells
quadrate to elongate, commonly sinuose or sinuose-nodulose, usually thick-walled. Perichaetia terminal
of stems or lateral
branches; perichaetial leaves differentiated or not. Seta short to long, smooth or rarely papillose
. Capsule usually erect, usually ovoid
, cylindrical or cupulate
or rarely strongly ventricose
at the base
, smooth or sulcate
; annulus present or absent, often compound
, deciduous or persistent
; operculum mammillate
to long-rostrate, sometimes attached to the columella after dehiscence (most Schistidium) ; peristome present, seldom rudimentary
or absent, consisting of 16 teeth, lanceolate to linear, entire, perforated or cribrose, variously split into 2 or 3 unequal prongs or divided
nearly to the base into two filiform
segments, smooth or variously ornamented. Calyptra small to large, covering only the operculum to half or more of the capsule, cucullate
, mitrate, or mitrate-campanulate, smooth or plicate, naked, sometimes papillose, slightly to distinctly lacerated or deeply lobed
at the base. Spores globose
, smooth or papillose.
Genera ca. 11, species ca. 325 (9 genera, 109 species in the flora ) : worldwide.
Most species in the Grimmiaceae are xerophytic and colonizers of bare, usually dry and exposed rocks and stones , forming predominantly dark green to blackish cushions or tufts. However, some species occur on wet or damp rocks along watercourses and lakes or in seepage sites. They rarely inhabit soil and only a few species are epiphytes.
The Grimmiaceae is classically distinguished by quadrate to short-rectangular mid leaf cells typically sinuose to nodulose and thick-walled, and leaves usually awned , often with the awns long and toothed or papillose. There is a wide range of variation . While awns are present in most species, length varies from a short, translucent apiculus to exceeding the length of the lamina. Awns can be flat or terete , smooth or denticulate , spinose or papillose, and long-decurrent or not. Similarly, mid leaf cells range from oblate to long-rectangular and sinuose to almost straight. The range of variation in these characters makes it difficult to describe this family in simple and unequivocal terms .
The generic classification within the Grimmiaceae has long been a subject of controversy. In traditional treatments the family is considered to include the two largest genera, Grimmia and Racomitrium, with several peripheral, mostly mono- or oligotypic genera, including Aligrimmia R. S. Williams, Coscinodon, Coscinodontella R. S. Williams, Indusiella, Jaffueliobryum, and Leucoperichaetium Magill. A number of segregates have been split from the large and heterogeneous Grimmia, namely Dryptodon Bridel, Guembelia Hampe, Hydrogrimmia (I. Hagen) Loeske, Orthogrimmia (Schimper) Ochyra & Zarnoweic, Schistidium, and Streptocolea I. Hagen. In this treatment, only Schistidium is accepted. Racomitrium has been divided into four sharply delimited genera and the group is recognized at the subfamily level.
R. Ochyra et al. (2003) used two peristome types to divide Grimmiaceae into two subfamilies, Grimmioideae and Racomitrioideae. The Schistidium-type peristome defines the Grimmioideae. This peristome has lanceolate teeth that are entire or perforate and distally usually split into two or three unequal prongs that are smooth to ornamented. They have a distinctly thick and trabeculate abaxial side and a thin adaxial side. There is no basal membrane . In the Racomitrium-type peristome, which defines the Racomitrioideae, the teeth are linear and divided nearly to the base into two filiform branches. The teeth are equally thickened and less prominently trabeculate on both adaxial and abaxial sides. They usually arise from a low, basal membrane and often have a prostome .
Plants (3-) 10-40(-180) mm, in dense cushions
, green, brown, or black, often with yellow, orange, or red tones. Leaves ovate-lanceolate, occasionally ovate-triangular, less commonly lanceolate to linear-lanceolate or elliptical
proximally, sharply keeled or nearly flat distally, margins
recurved, rarely plane
, distal lamina usually 1-stratose or 2-stratose in striae or patches, rarely 2-stratose, specialized laminal
chlorophyllose structures absent, muticous
to long-awned, sometimes ending in a fleshy
, multistratose apiculus
; basal cells
rectangular, with straight or sinuose and thin to thick cell walls
; mid leaf and distal cells quadrate
, rectangular, or ovate
, rarely sub-triangular, smooth
, usually sinuose and thick-walled. Gemmae absent. Sexual condition autoicous
, rarely dioicous; perichaetial leaves usually enlarged. Seta short, straight. Capsule erect
; annulus rudimentary
or absent; operculum rostrate
or rarely mamillate, usually falling attached to columella (except S. trichodon). Calyptra cucullate
or mitrate, not erose, not fully covering operculum, smooth.
Species ca. 120: North America, Mexico, Central America, South America, Eurasia , Africa, Atlantic Islands, Pacific Islands, Australia, Antarctica.
The genus Schistidium has consistently fascinated yet confounded bryologists across North America. Treatments vary from region to region, and names applied to specimens at both the species and varietal levels have been as inconsistent as the characters used to differentiate the taxa. The treatment of the S. apocarpum complex by H. H. Blom (1996) and a survey of Nordic species of Schistidium (Blom 1998) assist in a better understanding of the taxonomy of this complex genus in North America, but many problems of taxonomic interpretation remain. Although Schistidium offers a great number of both gametophytic and sporophytic characters for study, some traits are not well understood and further detailed field and laboratory research is needed.
When examining a specimen of Schistidium certain steps are helpful. It is important to examine the leaves proximal to the perichaetial region. Transverse-sections from the distal region to mid leaf of multiple leaves are also critical in most cases. The necessity of numerous transverse-sections is apparent when studying S. papillosum or S. boreale, for example, as some leaves can be slightly papillose and unless numerous sections are made the papillae may be missed. Mature , empty capsules that are not overly degraded, although not always available, should be used. Transverse-sections of the capsules assist in the easier examination of the exothecial cells and peristome teeth. It is also useful to examine more than one capsule if available, as there can be some variation in exothecial cell makeup. As H. H. Blom (1996) pointed out, mixed populations are present in some sites, especially in more humid areas, so care must be taken to ensure that all species in a collection are separated. Blom also provided a great amount of supplementary detail about many of the species treated here.
Species Schistidium apocarpum
Plants in open tufts or mats, olivaceous
to brownish (black),
sometimes with yellowish tones. Stems 1.2-12 cm, central strand
weak or absent. Leaves erect
, rarely (falcate-)
when dry, ovate-lanceolate, sharply keeled
1.7-2.5(-3.2) mm, 1-stratose or rarely 2-stratose in striae distally;
usually recurved throughout or to just before the apex, usually
distally, 1 or 2-stratose; apices acute or sub-obtuse;
or excurrent as a smooth
or weakly denticulate,
occasionally decurrent awn
surface often papillose
; distal laminal
cells mostly short-rectangular,
8-10 µm wide, smooth, sinuose. Sexual condition autoicous
Capsule dark red or brown, short-cylindric, 0.7-1.3 mm; exothecial
cells usually quadrate, sometimes mixed with short-elongate or oblate
cells, thin-walled, usually trigonous
; stomata present; peristome
or erect, often twisted, 350-600(-700) µm, red, densely
papillose, usually perforated. Spores 11-14(-19) µm,
or smooth. Capsules mature
to early summer.
Although H. H. Blom (1996) considered Schistidium apocarpum to be restricted in the flora area to the eastern portions of North America, it is more widespread and scattered across the continent. The denticulate leaf margins, papillose abaxial costal surface, short-cylindrical capsules, and the thin-walled, often evenly quadrate exothecial cells are distinguishing characters. The long peristome teeth are also useful in identifying S. apocarpum. In good condition the peristome can only be confused with that of S. trichodon, which differs in the darker, often black color of its plants , the peristome teeth often forming a dome, and the cylindrical capsules that retain the columella. [source]
Rocks in somewhat shaded habitats ; low to moderate elevations (0-1500 m ).
Typically found at an altitude of 0 to 4,701 meters (0 to 15,423 feet).
- Whittaker & Margulis,1978
- Haeckel, 1866
- Cavalier-Smith, 1981
- A. Braun, in Ascherson, 1860
- (Linnaeus, 1753) Cavalier-Smith, 1998
- Cavalier-Smith, 1998
- Bruch & Schimp., in Bruch et al., 1845, nom. cons.
- [Greek schistos, split or divided, and -idium, diminutive, alluding to peristome]
- Specific epithet:
- Bruch & W. P. Schimper in B.S.G., 1845
- Botanical name: - Schistidium apocarpum Bruch & W. P. Schimper in B.S.G., 1845 Bruch & W. P. Schimper in B.S.G., 1845
- Specific epithet: apocarpum - Bruch & W. P. Schimper in B.S.G., 1845
- Genus: Schistidium () - Bruch & Schimp., in Bruch et al., 1845, nom. cons. - [Greek schistos, split or divided, and -idium, diminutive, alluding to peristome]
- Family: Grimmiaceae () - Arn.
- Order: Grimmiales ()
- Superorder: Haplolepideae ()
- Subclass: Dicranidae ()
- Class: Bryopsida () - - Mosses
- Infraphylum: Bryatae () - Cavalier-Smith, 1998
- Subphylum: Musci () - (Linnaeus, 1753) Cavalier-Smith, 1998
- Phylum: Bryophyta () - A. Braun, in Ascherson, 1860 - Mosses
- Subkingdom: Viridaeplantae () - Cavalier-Smith, 1981
- Kingdom: Plantae () - Haeckel, 1866 - Plants
Grimmia apocarpa Hedwig • S. umbrosum (J. E. Zetterstedt) H. H. Blom • Schistidium lancifolium H. H. Blom
Status: Accepted Name
Last scrutiny: 19-Jul-2004
Members of the genus Schistidium
ZipcodeZoo has pages for 13 species, subspecies, varieties, forms, and cultivars in this genus:
S. agassizii (Agassiz's Schistidium Moss) · S. andreaeopsis (Schistidium Moss) · S. apocarpum (Schistidium Moss) · S. cinclidodonteum (Schistidium Moss) · S. cryptocarpum (Schistidium Moss) · S. heterophyllum (Schistidium Moss) · S. maritimum (Seaside Schistidium Moss) · S. obtusifolium (Obtuseleaf Schistidium Moss) · S. occidentale (Western Schistidium Moss) · S. pulvinatum (Pulvinate Schistidium Moss) · S. rivulare (Streamside Schistidium Moss) · S. tenerum (Schistidium Moss) · S. trichodon (Schistidium Moss)
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Accessed through GBIF Data Portal February 27, 2008:
- Australian Antarctic Data Centre: Australian Antarctic Division Herbarium
- Australian National Herbarium (CANB)
- Biologiezentrum der Oberoesterreichischen Landesmuseen: Biologiezentrum Linz
- Canadian Museum of Nature: Canadian Museum of Nature Herbarium
- European Environment Agency: EUNIS
- GBIF-Spain: Real Jardin Botanico, Madrid: MA-Musci
- GBIF-Spain: Universidad de Murcia, Dpto. Biología Vegetal (Botánica), Murcia: MUB-MUSCI
- GBIF-Sweden: Herbarium of Oskarshamn (OHN)
- GBIF-Sweden: Mosses (S)
- Icelandic Institute of Natural History: Herbarium (ICEL)
- Missouri Botanical Garden
- National Herbarium of New South Wales: NSW herbarium collection
- Natural History Museum, University of Oslo: Bryophyte herbarium, Bergen (BG)
- Natural History Museum, University of Oslo: Bryophyte herbarium, Oslo (O)
- The New York Botanical Garden: American Bryophyte Catalog
- UK National Biodiversity Network: British Bryological Society - Bryophyte data for Great Britain held by BRC for compilation
- UK National Biodiversity Network: Environment and Heritage Service - EHS Species Datasets
- UNIBIO, IBUNAM: MEXU/Colección de Briofitas
- University Museums of Norway (MUSIT)
- Biodiversity Heritage Library NamebankID: 3873407
- Catalogue of Life Accepted Name Code: MOS-35149157
- Global Biodiversity Information Facility Taxonkey: 4279666
- Integrated Taxonomic Information System (ITIS) Taxonomic Serial Number (TSN): 548090
- U.S.D.A. Plant Symbol: SCST9
- Zipcode Zoo Species Identifier: 39887
- Roxanne I. Hastings, Ryszard Ochyra "Grimmiaceae". in Flora of North America Vol. 27 Page 39, 204, 205, 231, 265, 266, 286, 294, 306, 615. Oxford University Press. Online at EFloras.org. [back]
- Terry T. McIntosh "Schistidium". in Flora of North America Vol. 27 Page 204, 205, 206, 207, 218. Oxford University Press. Online at EFloras.org. [back]
- "Schistidium apocarpum". in Flora of North America Vol. 27 Page 207, 210, 211, 217. Oxford University Press. Online at EFloras.org. [back]
- Mean = 256.760 meters (842.388 feet), Standard Deviation = 471.080 based on 65,620 observations. Altitude information for each observation from British Oceanographic Data Centre. [back]