BFNA Title: Cryphaeaceae
William D. Reese
Plants small to medium in size, mostly dull, with creeping primary stems bearing spreading to pendent branches, mostly brownish-green. Stems irregularly branched; paraphyllia lacking; pseudoparaphyllia lacking; axillary hairs 2--4 per axil, of 1--2 short proximal cells with brownish walls and 1--3 elongate distal cells. Branches short to elongate, simple to much-branched, terete. Leaves spirally inserted, imbricate, rapidly spreading when moistened, mostly ovate-acuminate, margins entire, or serrate distally, apex mostly acute; costa single; medial cells rhomboidal to fusiform, smooth to prorulose; alar cells rounded to quadrate. Sexual condition autoicous. Perigonia gemmiform, small, axillary. Perichaetia gemmiform, axillary or terminal on branches. Seta single, very short. Capsule immersed, erect, symmetric, stomates scanty, proximal; annulus usually revoluble; operculum conic-rostrate; peristome mostly double, pale; exostome of 16 teeth; endostome of 16 small linear segments or lacking, cilia lacking. Calyptra mitrate or subcucullate, smooth to papillose. Spores spherical, mostly granular to papillose, sometimes smooth.
Genera 8--10, species ca. 60--70 (2 genera, 5 species in the flora): worldwide, primarily tropical and subtropical regions.
The Cryphaeaceae are defined by the combination of creeping primary stems with firm spreading-erect branches, immersed capsule, corticolous habitat, dull aspect, and pale peristome. The vegetative leaves of many of the species of this family are very similar. The Cryphaeaceae as treated here comprise only Cryphaea and Schoenobryum. Several closely related genera in the family occur outside of the flora area (cf. H. Akiyama 1990; M. G. Manuel 1981). Other genera of the flora area, e.g., Alsia, Dendroalsia, and Forsstroemia, have been included in the concept of the Cryphaeaceae by various authors, e.g., M. R. Crosby and R. E. Magill (1981), but are here assigned to other families: Alsia and Dendroalsia in the Leucodontaceae and Forsstroemia in the Leptodontaceae.
SELECTED REFERENCES Akiyama,
H. 1990. Morphology and taxonomic significance of dormant branch primordia,
dormant buds, and vegetative reproductive organs in the suborders
Leucodontineae and Neckerineae (Musci, Isobryales). Bryologist 93: 395--408.
Crosby, M. R. and R. E. Magill. 1981. A Dictionary of Mosses. Monographs in
Systematic Botany from the
1. Perichaetia lateral; peristome double (single in C. ravenelii) . . . . 1. Cryphaea
1. Perichaetia terminal; peristome single . . . . 2. Schoenobryum
1. CRYPHAEA F. Weber, Tabula Calyptratarum Operculatarum, Musc. Frondos. Gen. 3. 1814 • Greek, cryph, hidden; referring to the immersed sporophytes
Plants small, dull, green to glaucous- or brownish-green. Branches mostly short, slender, wiry, simple or irregularly branched; axillary hairs 3--4 per axil, with 1(--2) short proximal cells and 2--3 elongate distal cells. Leaves tightly imbricate when dry, mostly wide-spreading when moist, concave, margins plane or somewhat recurved proximally; medial cells smooth or prorulose abaxially, especially distally, mostly smooth adaxially. Perichaetia lateral on branches; interior leaves mostly more or less awned, the awns usually denticulate-roughened. Peristome mostly double, exostome teeth narrowly triangular, papillose; endostome segments narrow, papillose, sometimes indistinct or lacking. Calyptra mostly conic, sometimes almost cucullate, smooth to papillose.
Species ca. 50--60 (4 in the flora): worldwide, mostly tropical and subtropical regions.
Plants of Cryphaea grow in thin or dense colonies, with the slender, often inconspicuous branches standing out from the substrate. The dull, wiry aspect of the spreading, mostly simple branches is characteristic.
SELECTED REFERENCE Manuel, M. G. 1973. Studies in Cryphaeaceae I. A revision of the genus Cryphaea in North America North of Mexico. Bryologist 76: 144--162.
1. Leaf apex rounded to bluntly acute; peristome single, exostome teeth joined in pairs . . . . 4. Cryphaea ravenelii
1. Leaf apex acute to acuminate; peristome double, exostome teeth single.
2. Costa ending in leaf acumen, percurrent or nearly so; leaf apices acuminate . . . . 3. Cryphaea nervosa
2. Costa ending at midleaf or above, but not percurrent; leaf apices acute.
3. Interior perichaetial leaves mostly 1.6--1.9 mm, their awns about 1/5--1/4 length of expanded portion of leaf; costa stout, often 2-fid at tip; plants common, widespread in eastern and southeastern U.S . . . . 2. Cryphaea glomerata
Interior perichaetial leaves mostly 2.5--2.8 mm, their awns about 1/2 length
of expanded portion of leaf; costa slender, not 2-fid at tip; plants rare, in
the flora area known only from southern
1. Cryphaea filiformis (Hedwig) Bridel, Musc. Recent. Suppl. 4: 139. 1819
Neckera filiformis Hedwig, Sp. Musc. 202. 1801; Cryphaea floridensis H. A. Crum; Cryphaea glomerata var. scabra Grout
Branches mostly elongate, often branched. Leaves narrowly acute to short-acuminate, wide-spreading when wet, mostly 1.3--1.6 mm; costa ending in base of acumen, often laterally spurred, not 2-fid at tip. Interior perichaetial leaves mostly 2.5--2.8 mm; awns distinct, strongly denticulate, about 1/2 length of expanded portion of leaf. Peristome double; exostome teeth single; endostome segments slenderly triangular. Calyptra conic. Spores papillose.
Capsules mature Mar.--Apr. Twigs and branches of trees and shrubs, in humid forests; 0 m; Fla.; Mexico; West Indies; Central America.
Vegetatively, C. filiformis
is rather similar to C. glomerata, but the branches of the former are
generally longer and the interior perichaetial leaves are much longer and
have proportionally longer awns. Although A. J. Grout (1934) wrote that the
capsules of C. filiformis are often at the tips of stems and branches,
this is not correct; he was misled by a mixed collection that included Schoenobryum
concavifolium in addition to C. filiformis. In S. concavifolium
the perichaetia are terminal. Cryphaea filiformis was attributed to
2. Cryphaea glomerata Sullivant in A. Gray, Man. N. U.S. Bot. ed. 2, 656. 1856
Branches short to elongate, mostly simple. Leaves broadly acute, wide-spreading when wet, mostly 0.8--1.2 mm; costa ending at midleaf or in base of acumen, often laterally spurred and often more or less 2-fid at tip. Interior perichaetial leaves mostly 1.6--1.9 mm; awn usually distinct, denticulate, about 1/5--1/4 length of expanded portion of leaf. Peristome double; exostome teeth single; endostome segments linear, slender. Calyptra broadly conic. Spores smooth or papillose.
Capsules mature Sept.--June. Twigs, branches, and trunks of shrubs and trees, on logs, rarely on rock, in humid forests; 0--1000 m; Ala., Ark., Conn., Del., Fla., Ga., Ky., La., Md., Mass., Miss., Mo., N.J., N.C., Ohio, Okla., S.C., Tenn., Tex., Va.
Cryphaea glomerata often grows mixed with C. nervosa,
but the two can be easily distinguished under low magnification by the more
narrowly pointed and plicate-appearing leaves of C. nervosa. The
ranges of C. glomerata and the subtropical C. filiformis
overlap in southern
3. Cryphaea nervosa (Hooker & Wilson) J. K. A. Müller, Linnaea 19: 211. 1846
Daltonia nervosa Hooker & Wilson in T. Drummond, Musci Amer. So. States 100. 1841
Branches short or elongate, mostly simple. Leaves acuminate, wide-spreading when wet, mostly 1.0--1.2 mm; costa percurrent or nearly so, not spurred, not 2-fid at tip. Interior perichaetial leaves mostly 1.6--1.9 mm; awn usually distinct, denticulate, about 1/5--1/4 length of expanded portion of leaf. Peristome double; exostome teeth single; endostome segments irregularly linear. Calyptra subcucullate to cucullate. Spores papillose.
Capsules mature Dec.--Apr. Twigs, branches, and trunks of trees in humid forests, sometimes in swamp forests; 0--1060 m; Ala., Fla., Ga., La., Miss., N.C., S.C., Tenn., Tex.
Plants of C. nervosa generally grow in more humid habitats than those of C. glomerata, a species with which its plants are often intermingled. The narrowly pointed leaves of C. nervosa make it easy to recognize in the field. The costa is often so prominent that the leaves of dry plants appear to be plicate.
Branches short to elongate, mostly simple. Leaves rounded to bluntly acute, ascending when wet, mostly 1 mm; costa ending in base of acumen, often laterally spurred and often 2-fid at tip. Interior perichaetial leaves mostly 1.6--1.8 mm, apiculate or with short rough awn to about 1/6--1/4 length of expanded portion of leaf. Peristome single; exostome teeth mostly joined in pairs; endostome lacking. Calyptra narrowly conic-subcucullate. Spores papillose.
Capsules mature Apr., May. Trunks
and branches of saplings and living and dead trees, exposed roots, forests;
to 330 m;
Cryphaea ravenelii is rare. Although similar at first glance to C. glomerata, it can be easily distinguished by its blunt leaf apices and paired exostome teeth. Further, the interior perichaetial leaves of C. glomerata have a greater tendency to be awned than those of C. ravenelii. Cryphaea ravenelii occupies the same habitats as C. glomerata and the two species have often been collected growing together.
2. SCHOENOBRYUM Dozy & Molkenboer, Musc. Frond. Ined. Archip. Indici fasc. 6: 183. 1848 • Greek, schoeno, a rope, and bryo, moss; referring to the perceived ropelike appearance of the stems, especially when dry
Plants small, dull, brownish-green, sometimes with reddish tinge. Branches mostly short, irregularly branched; axillary hairs 2--3 per axil, with 1 short proximal cell with brownish walls and 1--3 elongate distal cells. Leaves tightly imbricate when dry, spreading when moist, concave, margins plane to revolute; medial cells smooth to prorulose. Perichaetia terminal on branch tips, interior leaves awned. Peristome single, exostome teeth papillose. Calyptra conic, somewhat papillose distally.
Species ca. 24 (1 in the
flora): nearly worldwide (but not
SELECTED REFERENCE Manuel, M. G. 1977. Studies in Cryphaeaceae IV. New combinations in Schoenobryum. Bryologist 80: 522-524.
1. Schoenobryum concavifolium (
Branches elongate, usually branched. Leaves broadly acute to short-acuminate, wide-spreading when wet, mostly 1 mm, margins plane; costa sometimes laterally spurred, rarely 2-fid at tip. Interior perichaetial leaves mostly 2--2.3 mm, awns distinct, nearly smooth to denticulate, about 1/5 length of expanded portion of leaf. Exostome teeth narrowly triangular. Spores granular.
Capsules mature Apr. Twigs, branches, and tree trunks in humid forests; 0 m; Fla.; Mexico; West Indies; Central America; South America; Asia.
Schoenobryum concavifolium is very rare in the flora area, where
it is known only from a few specimens from
Editor's note: Rao (2001) has recently published a paper of relevance to the floral representation of this family.
Buck, W. R. 1994. Three additions to the United States moss flora. Bryologist 97: 89-90.
Grout, A. J. 1934. Cryphaeaceae. In: A. J. Grout, ed. Moss Flora of North America, North of Mexico. Vol. 3, pp. 222--226. Newfane, Vt. and New York.
Rao, P. 2001. Taxonomic studies on Cryphaea (Cryphaeaceae, Bryopsida). 3. Revision of European, African, Australian and Oceanian, and American species. Bryobrothera 7: 37--111.
Reese, W. D. 1995. Campylostelium saxicola new to Indiana, Cryphaea ravenelii new to Virginia, comments on Cryphaea filiformis and Schoenobryum concavifolium in Florida, a note on A. J. Grout's North American Musci Perfecti number 218, and an observation on O. E. Jenning's illustration for Ptychomitrium incurvum. Evansia 12: 157--160.
Sharp, A. J. H. Crum and P. M. Eckel. 1994. The Moss Flora of Mexico. 2 vols. New York.