BFNA Title: Stereophyllaceae
Author: W. Buck and R. Ireland
Date: October 9, 2010
Edit Level: R
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

BFNA Web site: http://www.mobot.org/plantscience/BFNA/bfnamenu.htm

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XX. STEREOPHYLLACEAE W. R. Buck & Ireland

 

William R. Buck and Robert R. Ireland

 

Plants medium to moderately large, in thin to dense, often glossy, flat mats. Stems prostrate, complanate to terete, simple or sparingly and irregularly branched; cortical cells small and thick-walled in cross section, or rarely with an outer layer of large thin-walled cells subtended by small thick-walled cells, internally with large, thin-walled cells; pseudoparaphyllia filamentous or less commonly foliose; rhizoids in clusters below leaf insertions, sometimes on base of costa or adjacent cells on dorsal surface of leaves, smooth or papillose; axillary hairs with a single brown basal cell and 2--6 hyaline apical cells. Leaves erect to wide-spreading, usually imbricate when dry, lanceolate to ovate, flat to somewhat concave, obtuse to long-acuminate, symmetric or rarely asymmetric, nondecurrent, margins plane or incurved, entire to serrulate beyond leaf middle; costa single, extending 1/3--3/4\x leaf length (rarely absent in some leaves); distal laminal cells linear to rhomboidal, thin- or thick-walled, smooth or rarely 1-papillose over the lumina or prorulose at distal ends on abaxial surface; alar cells differentiated, usually in large areas, quadrate to rectangular, often unequally distributed on either side of costa but always covering the adaxial surface of the costa, collenchymatous. Specialized asexual reproduction unknown. Sexual condition usually autoicous, sometimes dioicous. Seta solitary, straight, long, smooth. Capsule cernuous or erect, straight or slightly arcuate; exothecial cells thin- or thick-walled; annulus differentiated and deciduous, or undifferentiated; operculum conic to short-rostrate; peristome double, exostome teeth on outer surface cross-striolate or with papillae in horizontal rows below, papillose above, ± projecting on inner surface; endostome with a high or low basal membrane, the segments keeled, shorter than or almost as long as the teeth, cilia present or absent. Calyptra cucullate, smooth, naked. Spores spherical to ovoid, minutely papillose.

            Genera 8, about 28 species (3 genera, 3 species in the flora); occurring mostly at low elevations in subtropical and tropical regions; Florida, North Carolina and Texas.

 

selected references Buck, W. R. and R. R. Ireland. 1985. A reclassification of the Plagiotheciaceae. Nova Hedw. 41: 89--125. Grout, A. J. 1945. A revision of the North American species of Stereophyllum and Pilosium, with descriptions of some South American species. Bryologist 48: 60--70. Ireland, R. R. and W. R. Buck. 1994. Stereophyllaceae. Fl. Neotrop. Monogr. 65: 1--49.

 

1. Leaves similar throughout stems and branches; costa strong, extending 1/3--3/4\x leaf length; capsules cernuous or erect; exothecial cells not or scarcely collenchymatous; spores 9--27 \um .............................1. Stereophylloideae, p. XX

1. Leaves dimorphic, dorsal leaves symmetric, oblong-ovate, differentiated alar cells few, lateral leaves cultriform, oblong-lanceolate, differentiated alar cells numerous; costae weak, rarely reaching 1/3\x leaf length, present only on lateral leaves; capsules cernuous; exothecial cells collenchymatous; spores 7--9\um ……………………………………………............................2. Pilosioideae, p. XX

 

XXa. Stereophyllaceae subfam. Stereophylloideae M. Fleischer, Musci Fl. Buitenzorg 4: 1158. 1923 "Stereophylleae."

 

Stems complanate to terete, simple or sparingly and irregularly branched; pseudoparaphyllia filamentous or foliose; rhizoids smooth or papillose; axillary hairs with a single brown basal cell and 2--6 hyaline apical cells. Leaves similar throughout stems and branches, lanceolate to ovate, obtuse to long-acuminate, mostly symmetric; costa single, extending 1/3--3/4\x the leaf length; distal cells linear to rhomboidal, thin- or thick-walled, smooth or 1-papillose over the lumina or prorulose at upper ends on abaxial surfaces; alar cells differentiated, numerous, quadrate to rectangular, collenchymatous, mostly unequally distributed. Capsule cernuous or erect, straight or slightly arcuate; exothecial cells thin- or thick-walled; annulus present and deciduous, or absent; operculum conic to short-rostrate.

            Genera 7 (2 in the flora): tropical and south temperate.

 

1. Leaves abruptly acute to obtuse; leaf cells ± isodiametric, 1-papillose over lumina, sometimes smooth ..........................................................................1. Stereophyllum

1. Leaves acuminate; leaf cells elongate, smooth or minutely prorulose on abaxial surface .........................................................................................................2. Entodontopsis

 

1. STEREOPHYLLUM Mitten, J. Linn. Soc. Bot.. Suppl. 1: 117. 1859 * [Greek, stereos, thick and phyllos, leaf]

   Glossophyllum (Müller Hal.) Hampe; Hypnum sect. Glossophyllum Müller Hal.

 

Plants medium-sized, in loose to dense dull mats, complanate to subjulaceous, pale- to yellow-green, sometimes brown-tinged. Stems creeping, simple or sparingly and irregularly branched; cortical cells small, thick-walled, central strand lacking or poorly developed; pseudoparaphyllia filamentous; rhizoids smooth, in clusters below leaf insertions on abaxial surface of stems; axillary hairs with a single brown basal cell and 4 hyaline, elongate apical cells. Stem and branch leaves similar, erect-spreading when moist, usually imbricate and somewhat contorted when dry, complanate or somewhat concave, close, stiff, smooth, oblong-ovate to nearly lingulate, abruptly acute to obtuse, sometimes apiculate, symmetric, nondecurrent; margins plane, entire or serrulate near apex; costa single, extending 2/3--3/4\x the leaf length, stout, bulging on abaxial leaf surface; cells often thick-walled, distal to median cells rhomboidal, typically 1-papillose over cell lumina on both surfaces, sometimes smooth, walls not porose; alar regions distinctly differentiated, cells quadrate, rectangular or oblate, collenchymatous, extending across adaxial surface of costa. Sexual condition autoicous, often producing sporophytes; perigonia scattered along stems, bracts ovate, acute, costate, cells firm-walled, rhomboidal throughout, smooth; perichaetia at bases of stems, leaves oblong-lanceolate, serrulate, especially above, costate, cells thick-walled, linear-flexuose above, thin-walled and rectangular below, smooth. Seta smooth, elongate, straight to somewhat flexuose, orange to reddish brown. Capsule cernuous or rarely erect, straight or slightly arcuate, orange to brown, smooth, ellipsoidal to ovoid, contracted below mouth when dry; exothecial cells thin-walled; annulus of 2--3 rows of small, firm-walled cells, persistent; operculum short-rostrate, sometimes oblique; peristome double, exostome teeth bordered, shouldered, outer surface cross-striolate below, papillose above, inner surface projecting; endostome finely papillose, basal membrane high, segments keeled, perforate, almost as long as the teeth, cilia 1--3, shorter than the segments. Spores spherical to ovoid, papillose.

            Species 2 (1 in the flora): North, Central and South America, West Indies, Asia, Africa, Philippines, Australia.

 

1. Stereophyllum radiculosum (Hooker) Mitten, J. Linn. Soc., Bot. 12: 542. 1869

      Hookeria radiculosa Hooker, Musci Exot. 1: 51. 1818; Hypnum wrightii (Sullivant) Sullivant; Omalia wrightii Sullivant in A. Gray; Pterygophyllum indicum Bélanger; Stereophyllum indicum (Bélanger) Mitten; S. radiculosulum (Müller Hal.) A. Jaeger; S. wrightii (Sullivant) Renauld & Cardot

 

Stems to 3 cm, 1--2 mm wide. Leaves 1.0--2.5 x 0.4--1.2 mm; upper cells 14--38 x 7--9 \um; alar cells 9--19 x 9--14 \um, often many on one side of costa and few on the other. Seta 0.6--1.2 cm. Capsule 0.7--1.8 mm; operculum 0.5--0.7 mm. Spores 12--24 \um.

            Oak woods or occasionally swamps, bases of trees, exposed roots, stumps, logs or sometimes on limestone; 0--100 m; Ala., Fla., Tex.; Mexico; West Indies; Central America; South America; Africa; Asia; Australia (Queensland).

            Stereophyllum radiculosum is immediately recognized by the dull, complanate plants with simple or sparingly branched stems, by singly costate leaves with a prominent costa that bulges on the dorsal surface, by abruptly acute to obtuse leaf apices, and by short, rhomboidal leaf cells, usually with a single papilla over the lumina on both surfaces of the leaves.

 

2. ENTODONTOPSIS Brotherus in H. G. A. Engler & K. Prantl, Nat. Pflanzenfam. 1(3): 895. 1907 * [Greek, entos, inside, odon, tooth, opsis, appearance]

   Hypnum subgen. Complanato-hypnum Hampe

 

Plants medium-sized, in thin to dense glossy mats, complanate to julaceous, light green to yellowish or brownish green. Stems creeping, simple or sparingly and irregularly branched; cortical cells small, thick-walled, central strand lacking or poorly developed; rhizoids smooth, in clusters below leaf insertions on abaxial surface of stems; pseudoparaphyllia filamentous; axillary hairs with a single brown basal cell and 3--6 hyaline apical cells. Stem and branch leaves similar, stiff to lax, close to distant, erect spreading when moist, usually imbricate and somewhat contorted when dry, sometimes homomallous toward substrate, flat to concave, smooth, rarely wrinkled, oblong-lanceolate, oblong-ovate, ovate-lanceolate or sometimes lingulate, acuminate, acute or obtuse, symmetric or asymmetric, nondecurrent; margins plane or incurved below middle, serrate to serrulate distally, serrulate to entire proximally, sometimes entire throughout; costa single, 1/3--1/2\x leaf length; cells thin-walled, distal cells broadly fusiform with tapering ends, sometimes with short, rhomboidal distal cells in obtuse leaves, mostly smooth, occasionally prorulose at upper ends on abaxial surface, walls not porose; alar regions strongly differentiated, cells quadrate, rectangular or oblate, large, often many on one side of costa and few on other, collenchymatous, extending over adaxial surface of costa. Sexual condition autoicous, often producing sporophytes; perigonia scattered along stems, bracts ovate, ecostate; perichaetia at bases of stems, leaves ovate, acuminate, entire to serrulate, cells linear-rhomboidal above, laxly rectangular proximally, costate to ecostate within a single perichaetium. Seta smooth, elongate, straight to somewhat flexuose, orange to reddish brown. Capsule erect, inclined or cernuous, straight or slightly arcuate, orange to brown, smooth, cylindric, ellipsoidal or ovoid, often contracted below mouth when dry; exothecial cells thin-walled; annulus of 2--3 rows of firm-walled cells, often falling with the operculum; operculum conic to short-rostrate; peristome double, exostome teeth bordered, shouldered, outer surface cross-striolate basally, papillose distally, inner surface strongly to scarcely projecting; endostome smooth or papillose, basal membrane high to low, segments keeled, perforate, somewhat shorter than the teeth, cilia 1--3, sometimes rudimentary, shorter than the segments. Spores spherical to ovoid, papillose.

            Species ca. 17 (1 in the flora): North, Central and South America, West Indies, Asia, Africa.

 

1. Entodontopsis leucostega (Bridel) W. R. Buck & Ireland, Nova Hedw. 41: 103. 1985

 

   Leskea leucostega Bridel, Bryol. Univ. 2: 333. 1827; Stereophyllum leucostegum   (Bridel) Mitten

 

Plants in loose or dense, glossy mats, complanate-foliate to subjulaceous, pale- to yellow-green, brownish green with age. Stems to 3 cm, 1--3 mm wide; axillary hairs with a single short brown basal cell and 3 elongate hyaline apical cells. Stem and branch leaves 1--2 x 0.4--0.8 mm, flat or sometimes slightly concave, ovate-lanceolate to oblong-ovate, acuminate, rarely acute; margins plane or incurved below the middle, entire or serrulate near the apex; costa 1/2--3/4\x leaf length; cells smooth or minutely prorulose on abaxial surfaces, median cells 47--165 x 7--9 \um; alar cells quadrate to short-rectangular, sometimes oblate, 14--33 x 11--24 \um. Seta yellowish to reddish brown, 0.5--0.8 cm. Capsule orange to reddish brown, 0.8--1.5 mm, contracted below mouth when dry; operculum obliquely conic to short-rostrate, 0.2--0.5 mm. Spores 15--27 \um.

            Woods, rotten logs, exposed tree roots; 0--100 m; Fla.; Mexico; West Indies; Central America; South America; Asia; Africa.

            Entodontopsis leucostega is recognized by the glossy, complanate-foliate plants with erect-spreading, narrow, acuminate leaves, by the elongate laminal cells with tapering ends that are smooth or sometimes prorulose on the abaxial surface and by the erect or inclined to cernuous capsules. A. J. Grout (1945) revised the North and Central American species, placing them in the genus Stereophyllum.

 

XXb. Stereophyllaceae subfam. Pilosioideae W. R. Buck & Ireland, Fl. Neotrop. Monogr. 65: 42. 1994

 

Stems complanate, irregularly branched; pseudoparaphyllia filamentous; rhizoids smooth; axillary hairs with a single brown basal cell and 3 hyaline apical cells. Leaves

dimorphic, obtuse to short-acute, abaxial leaves oblong-ovate, symmetric, lateral leaves oblong-lanceolate, cultriform; costa single in lateral leaves, weak, lacking in dorsal leaves; distal cells linear, thin-walled, smooth; alar cells differentiated in lateral leaves, numerous, large, oblong, colored, abaxial leaves with few differentiated cells, small, quadrate to subquadrate. Capsules cernuous, symmetric; exothecial cells collenchymatous; annulus undifferentiated; operculum short-rostrate.

            Genus 1: Neotropical.

 

1. PILOSIUM (Müller Hal.) M. Fleischer, Musci Fl. Buitenzorg 4: 1158. 1923 * [Greek, pilos, cap; Latin, ium, quality or nature of]

   Stereophyllum sect. Pilosium Müller Hal., Flora 83: 340. 1897

 

Plants medium-sized, in dense glossy mats, complanate, yellowish green. Stems creeping, irregularly branched; cortical cells small, thick-walled, central strand lacking; rhizoids in clusters below leaf insertions on abaxial surface of stems and branches; pseudoparaphyllia filamentous. Leaves dimorphic, somewhat stiff, close, wide-spreading, somewhat curled when dry, curled downward when moist, flat to slightly concave, non-plicate, obtuse to short-acute, non-decurrent, abaxial leaves oblong-ovate, symmetric, lateral leaves oblong-lanceolate, cultriform; margins plane, entire to serrulate, especially distally; costa single in lateral leaves, weak, rarely extending 1/3\x length of leaf, lacking in abaxial leaves; cells thin-walled, distal and median cells linear, smooth, heavily chlorophyllose, non-porose; alar cells differentiated in lateral leaves, numerous, large, oblong, colored, porose, extending across adaxial surface of costa, abaxial leaves with few differentiated cells, small, quadrate to subquadrate. Sexual condition autoicous, often producing sporophytes; perigonia scattered along stems, bracts ovate, ecostate; perichaetia scattered along stems, often in pairs, leaves ovate to lanceolate, deeply erose, ecostate, cells linear. Seta smooth, long, straight, orange-red. Capsule cernuous, symmetric, orange-red, smooth, short-cylindric, contracted under mouth when dry; exothecial cells collenchymatous; annulus undifferentiated; operculum short-rostrate; peristome double, exostome teeth bordered, outer surface cross-striolate below, papillose above, inner surface projecting; endostome papillose, basal membrane high, segments keeled, perforate, shorter than the teeth, cilia single, shorter than the segments. Spores spherical, papillose.

            Species 1: North, Central and South America, West Indies.

 

1. Pilosium chlorophyllum (Hornschuch) Müller Hal., Flora 83: 340. 1897

   Hypnum chlorophyllum Hornschuch, Fl. Bras. 1(2): 89. 1840; Stereophyllum chlorophyllum (Hornschuch) Mitten

 

Plants in dense, glossy mats, complanate, yellowish green. Stems to 6 cm, 2--4 mm wide, irregularly branched. Leaves dimorphic, abaxial leaves 1.3--1.6 x 0.6--0.9 mm, oblong-ovate, symmetric, lateral leaves 1.5--2 x 0.6--1 mm, oblong-lanceolate, cultriform; margins plane, entire to serrulate, especially distally; costa single in lateral leaves, extending a short distance above base, lacking in abaxial leaves; median cells 89--146 x 7--12 \um; alar cells strongly differentiated in lateral leaves, colored, oblong, 24--89 x 14--28 \um, weakly differentiated in abaxial leaves, quadrate to subquadrate, 12--24 x 9--12 \um. Seta 1.5--1.8 cm. Capsule 0.8--1.2 mm; operculum 0.4--0.6 mm. Spores 7--9 \um.

            Forested sheltered gorge, rotten logs, exposed tree roots; 635--730 m; known from a single collection in our range, NC.; Mexico; West Indies; Central America; northern South America.

            Pilosium chlorophyllum is rarely confused with the other species in the Stereophyllaceae. It is distinctive because of its glossy, complanate plants that have dimorphic leaves; the abaxial ones symmetric, oblong-ovate, with neither a costa nor a distinctive alar region of differentiated cells; and the lateral ones cultriform, oblong-lanceolate, with a short, weak costa scarcely reaching 1/3\x leaf length, and a large area of enlarged, colored alar cells on one side of the costa. The cernuous, short-cylindric capsules have collenchymatous exothecial cells and the spores, 7--9 \um in diameter, are smaller than the other members of the family.