BFNA Title: Taxiphyllum
Author: R. R. Ireland
Date: July 2, 2004
Edit Level: R 
Version: 1
Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

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Taxiphyllum - Hypnaceae

 

XX.   TAXIPHYLLUM  Fleischer, Musci Fl. Buitenzorg 4: 1434. 1922 * [Greek, tax, arrangement and phyll, leaf]

Robert R. Ireland

Plagiothecium subg. Taxiphyllum (M. Fleischer) Grout

 

Plants medium- to large-sized, in thin to dense mats, complanate-foliate, sometimes julaceous or subjulaceous, glossy, light- to dark-green or yellow-green.  Stems creeping, simple or sparingly and irregularly branched, cortical cells small, thick-walled, central strand sometimes present; rhizoids smooth, in clusters below leaf insertions usually on ventral surface of stems; axillary hairs with 1--2(--3) short brown basal cells and a single elongate apical cell; pseudoparaphyllia large to small, foliose.  Stem and branch leaves similar, stiff to flaccid, crowded and imbricate to distant, erect to wide-spreading or squarrose, often complanate, rarely secund at the tips, smooth or occasionally plicate, plane to concave, symmetric or nearly so, nondecurrent, ovate, ovate-lanceolate, oblong-lanceolate, oblong-ovate or linear-oblong, acute to subobtuse, sometimes acuminate; margins plane or recurved, serrate to serrulate above leaf middle, serrulate to entire below; costa short and double, sometimes lacking; cells moderately thick-walled, linear-flexuose, rhomboidal near leaf apex, smooth or prorate at distal ends on abaxial leaf surface, with walls not pitted; alar regions distinctly differentiated with 1 to several rows of quadrate to rectangular cells, or undifferentiated or nearly so.  Specialized asexual reproduction lacking.  Sexual condition dioicous or rarely synoicous, usually sterile.   Perigonia and perichaetia at base of stems and branches, perigonial bracts small, ovate, perichaetial bracts lanceolate to ovate, slenderly acuminate, ± spreading from an erect base, margins plane.  Seta smooth, elongate, straight or somewhat flexuose, dark red to brown.  Capsule inclined to cernuous, straight to subarcuate, yellowish brown to reddish brown, smooth, oblong-ovoid, somewhat contracted below mouth and wrinkled at neck when dry; opercula obliquely rostrate, often shorter than urn; annulus present, persistent, of 2 rows of cells; peristome cross-striolate below, papillose distally, bordered, trabeculate; endostome with a high to low basal membrane, keeled segments and cilia shorter than or approximately the same length as the segments, in groups of 1--3. Calyptra cucullate, smooth, naked.  Spores globose to ovoid, smooth to minutely papillose, 7--28 µm.

 

Species10--15 (4 in the flora): terrestrial habitats, primarily on calcareous substrates, temperate, subtropical or tropical regions; North, Central and South America, West Indies, Europe, Asia, Africa, Australia.

 

SELECTED REFERENCES  Crum, H. A. and L. E. Anderson.  1981.  Mosses of Eastern North America.  Vol. 2.  New York.  Ireland, R. R.  1982.  Moss Flora of the Maritime Provinces.  National Museums of Canada, Natl. Mus. Nat. Sciences, Publs. in Botany, No. 13. Ottawa.  Ireland, R. R.  1969.  A taxonomic revision of the genus Plagiothecium for North America, north of Mexico.  National Museums of  Canada, Natl. Mus. Nat. Sciences, Publs.in Botany 1: 1--118.  Ireland, R. R.  1986.  Synopsis of the genus Taxiphyllum for North America.  Lindbergia 12: 153--157.  Iwatsuki, Z.  1963.  Bryological Miscellanies XII.  Preliminary notes on the Japanese species of Taxiphyllum and its related genera.  J. Hattori Bot. Lab. 26: 63--69. 

 

1.  Plants julaceous to subjulaceous; leaves acuminate; alar cells noticeably differentiated, quadrate to short-rectangular in 1--several rows and extending 5--15 cells up the margins........... l. Taxiphyllum cuspidifolium

1.  Plants complanate-foliate; leaves acute to acuminate, rarely subobtuse; alar cells not differentiated or quadrate to short-rectangular in 1--several rows and extending 1--9 cells up the margins.

2.  Leaves close, appressed-imbricate, never squarrose; margins usually plane or rarely recurved at base; quadrate alar cells numerous, in 1--several rows and extending 3--9 cells up the margins       3. Taxiphyllum deplanatum 

2.  Leaves usually remote or if imbricate not appressed, often squarrose; margins usually recurved to leaf middle; quadrate alar cells sparse, in 1--3 rows and extending 1--5 cells up the margins.

3.  Leaves broadly ovate-lanceolate, 0.8--1.6 mm wide; apices often acuminate and twisted; margins plane, rarely recurved; leaf cells smooth...... 2. Taxiphyllum alternans

3.  Leaves narrowly oblong-lanceolate or ovate-lanceolate, 0.3--0.6 mm wide; apices acute to acuminate, rarely subobtuse, not twisted; margins usually narrowly and indistinctly recurved to leaf middle; leaf cells sometimes dorsally prorate above    4. Taxiphyllum taxirameum

 

l.  Taxiphyllum cuspidifolium (Cardot) Z. Iwatsuki, J. Hattori Bot. Lab. 28: 220.  1965

 

Isopterygium cuspidifolium Cardot, Bull. Soc. Bot. Genève 4, sér. 2: 387.  1912 Plagiothecium mariannae A. J. Grout; Taxiphyllum mariannae (A. J. Grout) Schornherst 

 

Plants in thin mats, dark- to yellowish-green, with an oily sheen when wet.  Stems to 3 cm, 1--3 mm wide, prostrate, often radiculose ventrally.  Leaves loosely imbricate, usually concave, smooth, symmetric, 1.0--2.5 × 0.5--1 mm, ovate to broadly ovate-lanceolate, acuminate or filiform-acuminate, often twisted at apex, margins plane, serrulate to serrate beyond leaf middle, serrulate to entire proximally; costa short and double, one branch extending 1/3--1/2 length of leaf, rarely lacking; cells smooth; median cells 75--120 × 7--12 µm; alar cells 12--48 × 10--22 µm, quadrate to rectangular, in 2--several rows with 5--12 cells in marginal row.  Sexual condition dioicous.  Perichaetia large, numerous, bracts lanceolate to ovate-lanceolate, acuminate to sharply acuminate.   Perigonia and sporophytes unknown in North America. 

 

Calcareous soil and rock, rarely over exposed tree roots; 0--300 m; Ala., Fla., Tenn.; Asia.

 

Taxiphyllum cuspidifolium is distinguished by its julaceous to subjulaceous plants with an oily sheen when wet and by its loosely imbricate, erect-spreading, concave leaves with plane margins and acuminate to filiform-acuminate apices.  It is rare in North America where it is known from only one locality in Alabama (Bibb Co.), four in Florida (Alachua, Citrus, Jackson and Walton counties) and two in Tennessee (Anderson and Montgomery counties).  It has often been confused with Plagiothecium cavifolium (Bridel) Z. Iwatsuki, which has a more northern distribution.  For microscopic differences between the genera Plagiothecium and Taxiphyllum see the discussion under T. alternans.

 

2.  Taxiphyllum alternans (Cardot) Z. Iwatsuki, J. Hattori Bot. Lab. 26: 67.  1963

 

Isopterygium alternans Cardot, Beih. Bot. Centralbl. 17: 37.  1904 

 

Plants in thin mats, light- to yellowish-green or yellowish brown, glossy.  Stems to 6 cm,  3--5 mm wide, prostrate, naked or sometimes radiculose ventrally.  Leaves distant, becoming close and loosely imbricate at stem apices, spreading wet or dry, complanate, smooth, symmetric or asymmetric, 1.5--3.5 × 0.8--1.6 mm, ovate to broadly ovate-lanceolate, acuminate to filiform-acuminate, frequently twisted at apex, margins plane or rarely narrowly recurved for a short distance at base, serrulate to serrate beyond leaf middle, serrulate to entire proximally; costa short and double, one branch often extending 1/3--1/2 length of leaf, rarely lacking; cells smooth; median cells 84--156 × 9--12 µm; alar cells 14--43 × 14--20 µm, quadrate to long-rectangular, in 1--3 rows, seldom more, with 2--5 cells in marginal row.  Sexual condition dioicous.  Perichaetia large, numerous, bracts lanceolate to ovate-lanceolate, acuminate.  Perigonia and sporophytes  unknown in North America.

 

Soil, humus, exposed tree roots, rotten wood and logs in swamps, rarely on calcareous boulders beside streams and waterfalls; 0--500 m; Ala., Fla., La., Md., N.C., S.C.; Asia.   

 

Taxiphyllum alternans, like the previous species, is rare in North America.  It is recognized by distant leaves that are ovate to broadly ovate-lanceolate with acuminate to filiform-acuminate, frequently twisted apices, plane margins and with 2–5 quadrate to short-rectangular cells in the marginal rows.  It occurs from only a few localities in the southeastern United States with disjunct occurrences in Maryland (Charles and Montgomery Counties).  It too has sometimes been confused with Plagiothecium cavifolium (Bridel) Z. Iwatsuki but all Taxiphyllum species differ from Plagiothecium microscopically by their nondecurrent leaves and the presence of foliose pseudoparaphyllia.

 

3.  Taxiphyllum deplanatum (Sullivant) M. Fleischer, Musci Fl. Buitenzorg 4: 1435. 1923

 

Hypnum deplanatum Sullivant, Manual: 670. 1848; Isopterygium deplanatum (Sullivant) Mitten; Plagiothecium deplanatum (Sullivant) Spruce; Rhynchostegium deplanatum (Sullivant) Rau & Hervey

 

Plants in thin to dense mats, light- to yellow-green, glossy.  Stems to 4 cm, 1--3 mm wide, complanate, radiculose ventrally.  Leaves flaccid, appressed-imbricate, somewhat concave or flat, smooth, symmetric to somewhat asymmetric, 0.9--2 × 0.4--0.8 mm, ovate to ovate-lanceolate, rarely oblong-lanceolate or oblong-ovate, acuminate or often abruptly narrowed to an acute or filiform apex, margins plane or narrowly recurved for a short distance at base, serrulate to strongly serrate beyond leaf middle, serrate to serrulate proximally; costa very short and double or lacking; cells smooth; median cells 47--136 × 7--12 µm; alar cells 12--27 × 9--17 µm, quadrate to short-rectangular, in 1--several rows with 3--8 cells in the marginal row.  Sexual condition dioicous.   Perigonia unknown.  Perichaetia small, numerous, the bracts lanceolatae to ovate-lanceolate, acuminate.  Seta yellowish brown to red, flexuose, 0.7--1 cm.  Capsule yellowish brown to light brown, cernuous, 0.8--1.5 × 0.3--0.5 mm, oblong or ovoid, straight or arcuate, when dry contracted below mouth and tapered to a wrinkled neck; opercula obliquely rostrate, 0.3--0.5 mm.  Spores 11--13 µm. 

 

Capsules extremely rare but evidently mature in fall.  Shaded siliceous or calcareous soil and rock, often on bases of trees, exposed tree roots, and rotten logs, sometimes in cedar swamps; 60--2700 m; Man., N.B., Ont., Que., Sask.; Ariz., Ark., Conn., D.C., Ga., Ill., Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Nebr., N.H., N.J., N.Mex., N.Y., N.C., Ohio, Okla., Pa., S.Dak., Tenn., Vt., W.Va., Va., Wis.; Mexico; Central America (Honduras).

 

 

Taxiphyllum deplanatum  is widespread in northeastern North America with disjunct populations in Saskatchewan, Quebec, New Brunswick, Arizona and New Mexico.  Taxiphyllum deplanatum has often been confused with T. taxirameum but the two are easily distinguished superficially and microscopically.   The flaccid, appressed-imbricate leaves  characteristic of T. deplanatum will easily separate the species from T. taxirameum with its rigid, usually distant, wide-spreading to squarrose leaves.   The alar regions of T. deplanatum are well differentiated with quadrate to short-rectangular cells, 3--8 in the marginal rows, which is in striking contrast to the poorly differentiated alar regions of T. taxirameum that have only a few rectangular cells on the margins. Taxiphyllum deplanatum has been reported for the Gulf Coast region, based on a Louisiana specimen (R. R. Ireland 1969), however, it was later re-examined and found to be a misidentified specimen of T. taxirameum as W. R. Reese (1984) noted.

 

4.  Taxiphyllum taxirameum (Mitten) M. Fleischer, Musci Fl. Buitenzorg 4: 1435. 1922

 

Stereodon taxirameus Mitten, J. Proc. Linn. Soc. Bot. Suppl. 1: 105. 1859; Isopterygium geophilum (Austin) A. Jaeger; I. taxirameum (Mitten) A. Jaeger; Plagiothecium geophilum (Austin) Grout; Rhynchostegium geophilum Austin;  Taxiphyllum geophilum (Austin) Fleischer

 

Plants in thin to dense mats, dark- to yellow-green, glossy.  Stems to 6 cm, 2--4 mm wide, prostrate, rarely radiculose ventrally.  Leaves rigid, usually distant, wide-spreading to squarrose, flat or somewhat concave, smooth or plicate, symmetric or nearly so, 1--2 × 0.3--0.6 mm, ovate- or oblong-lanceolate, rarely narrowly ovate, acuminate or abruptly narrowed to an acute or rarely subobtuse apex, margins very narrowly recurved almost to apex, sometimes plane, serrulate to serrate throughout; costa lacking or short and double; cells smooth or prorate at distal ends on abaxial leaf surface; median cells 66--120 × 3--7 µm; alar cells 37--63 × 5--9 µm, long- to short-rectangular or rarely quadrate, in 1--3 rows with 1--5 cells in the marginal rows.  Sexual condition dioicous.  Perigonia small, bracts ovate, acute; Perichaetia small, bracts lanceolate to ovate-lanceolate, acuminate.  Seta reddish brown, flexuose, 0.7--1.2 cm.  Capsule reddish brown, cernuous, 1--1.5 mm, oblong-ovoid, arcuate, when dry slightly contracted below mouth and wrinkled at neck; opercula obliquely long-rostrate, 0.5--1 mm.  Spores 11--13 µm.

 

In shade, siliceous or calcareous soil and rock; 40--1500 m; Ala., Ariz., Ark., Del., D.C., Fla., Ga., Ill., Ind., Iowa, Kans., Ky., La., Md., Miss., Mo., N.J., N.Mex., N.Y., N.C., Ohio, Okla., Pa., S.C., Tenn., Tex., Va., W.Va.; Mexico; West Indies; Central and South America; Asia; East Indies.

 

These plants are typically glossy, complanate-foliate with the leaves stiff, distant, seldom overlapping, wide-spreading to squarrose, ovate-lanceolate or oblong-lanceolate with acute to subobtuse apices, narrowly recurved margins nearly to apex and poorly differentiated alar cells in 1--3 rows with 1--5 short-rectangular cells in the marginal rows. This is a polymorphic species common in the southeastern and south-central United States, particularly the Ozark Mountain region.  It has often been confused with the more northern T. deplanatum; for distinctions see the discussion under that species. Taxiphyllum taxirameum is the most common species of Taxiphyllum in Latin America and it is especially prevalent in parts of Mexico and the West Indies.

 

OTHER REFERENCES 

 

Ireland, R. R.  1994.  Taxiphyllum.  In: A. J. Sharp, H. Crum and P. M. Eckel(eds.), The Moss Flora of Mexico, Part 2. Orthotrichales to Polytrichales.  Mem. New York Bot. Gard. 69: 1022--1025. 

 

Mishler, B. D. & N. G. Miller.  1983.  Distributional studies of Massachusetts bryophytes. Rhodora 85: 421--432. 

 

Reese, W. D.  1984.  Mosses of the Gulf South.  Baton Rouge.