BFNA Title: Barbilophozia
Date: 25 Nov. 2020
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Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
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XX. BARBILOPHOZIA Loeske, Verh. Bot. Vereins Prov. Brandenburg 49(1): 37. 1907 * [Greek barbil, bearded, alluding to cilia on the leaves, and lophos, crest, spike, alluding to the angular ends of leaf lobes]

Terry T. McIntosh

Richard H. Zander

Alexey D. Potemkin

 

Neoorthocaulis L. Söderström, De Roo & Hedderson; Schljakovia Konstant & Vilnet; Schljakovianthus Konstant & Vilnet

 

Plants small to medium-sized, in loose to dense turfs or mats, prostrate, ascending, sub-erect, or erect, green, yellow-green, grayish-green, or dark brown, sometimes with a reddish or purple color. Stems branching terminal or lateral intercalary or occasionally both, transverse section with cortex 1--2(--3)-stratose; rhizoids sparse to common, colorless or pigmented proximally. Leaves succubous, obliquely, sometimes nearly transversely, inserted, distant to imbricate, spreading to appressed, flat to concave, obtrapezoidal, sub-orbicular, sub-quadrate or sub-rectangular, reniform, obcuneate-quadrate, rarely nearly symmetric, divided by variously shaped sinuses into 2--4 lobes; lobes entire or, sometimes lightly toothed; lobes entire or sometimes apiculate or mucronate; sinuses of various lengths, gibbous ot not; central laminal cells usually thin-walled and often trigonous, rarely papillose; oil bodies 2--25 per cell, biconcentric or not. Underleaves usually present, usually 2-lobed, margins often ciliate. Specialized asexual reproduction by gemmae at apices of leaf-lobes or shoots, or absent; gemmae 1--2-celled, smooth or angular. Sexual condition dioicous. Androecia intercalary, 1--4-androus, antheridial stalk 1--2-seriate. Gynoecia terminal, female bracts in 1 pair, usually 3--6-lobed. Perianth cylindric, mouth somewhat lobed and usually dentate or dentate ciliate, weakly to strongly plicate; perigynium absent. Capsule ovoid, walls 3--5-stratose, exterior wall with nodular thickenings, interior wall with semi-annular U-shaped thickenings, reddish brown. Elaters 2-spiraled, reddish brown. Spores spheric, 10--18 \um, papillose, brown.

 

Species 14 (12 in the flora): North America, South America, Eurasia, Africa, Australia, Oceania, Antarctica.

 

Barbilophozia and Anastrophyllum are both here recognized in Scapaniaceae. Both genera were moved to a new family Anastrophyllaceae L. Söderström, De Roo & Hedderson (L. Söderström  et al. 2010), which was distinguished in part with molecular traits. They pointed out, however, that “the morphological characters that single out the family are not yet clear.” They also proposed the genus Neoorthocaulis L. Söderström, De Roo & Hedderson for four species of Barbilophozia largely based on molecular evidence. New genera, Schljiakovia Konstantinova & Vilnet and Schljiakovianthus Konstantinova & Vilnet, which included species of Barbilophozia, were also published by N. A. Konstantinova and A. A. Vilnet (2009) on a molecular basis. These classifications are here considered presently impractical, and a classical approach is adopted.

 

Diminutive plants of Barbilophozia are often difficult or impossible to separate from some species of Lophozia.

 

SELECTED REFERENCES  Hong W. S. 2002. The distribution of Lophozia in North America west of the hundredth meridian. Lindbergia. 27: 49--62. Konstantinova, N. A. and A. A. Vilnet. 2009 [2010]. New taxa and new combinations in Jungermanniales (Hepaticae). Arctoa 18: 65--67. Schuster R. M. 1969. Lophozia (Dumort.) Dumort. In: R. M. Schuster. 1969. Hepaticae and Anthocerotae of North America. New York and London. Vol. 2.  Schuster, R. M. and K. Damsholt. 1987. 1. New taxa from South Greenland. Phytologia 63(5): 325--328.  Söderström, L., R. De Roo and T. Hedderson. 2010. Taxonomic novelties resulting from recent reclassification of the Lophoziaceae/Scapaniaceae clade. Phytotaxa 3: 47--53.  Stotler, R. E. and B. Crandall-Stotler. 2017. A synopsis of the liverwort flora of North America north of Mexico. Annals Missouri Bot. Gard. 102: 574--709.

 

1. Lateral leaves mostly 2-lobed.

2. Lateral leaves evenly concave and nearly hemispheric, lobes not gibbous; underleaves usually absent, if present minute……………………. 5. Barbilophozia cavifolia

2. Lateral leaves never evenly concave nor hemispheric; lobes +/- gibbous; underleaves present and distinct, lanceolate or 2-lobed.

3. Plants small, ca. 0.3--0.6 mm wide, blackish to olive-brown; leaves subtransversely inserted; laminal cells usually coarsely papillose with large, low papillae; Arctic …………… 11. Barbilophozia quadriloba (in part)

3. Plants larger, ca. 0.6--1 mm wide, green and yellow brown to brown-fuscous; leaves mostly obliquely inserted; laminal cells lacking large, low papillae.

4. Plants light green and yellow brown; leaves with broadly ovate to triangular ± concave to plane lobes; oil bodies (1--)2--4(--5) per cell; Arcto-boreal-montane ………………………….…. 9. Barbilophozia kunzeana (in part)

4. Plants green to brown and brown-fuscous; Leaves with broadly triangular acute ± concave lobes; oil bodies (4--)7--15 per cell; Arctic only………………………………. 8. Barbilophozia hyperborea (in part)

1. Lateral leaves mostly (2--)3--4-lobed.

5. Lateral leaves subtransversely inserted and oriented, deeply lobed with sinuses 2/5--3/5 leaf length, lobes usually 4, but some plants strictly 3-lobed; laminal cells coarsely papillose with numerous large, low papillae           11. Barbilophozia quadriloba (in part)

 5.  Lateral leaves obliquely inserted and oriented, more shallowly lobed with sinuses 1/5- 2/5 leaf length or less, lobes usually 3 or 4; laminal cells never coarsely papillose.

6. Leaves mostly (3--)4-lobed.

7........... Lateral leaves only slightly narrowed to the base, nearly symmetric, lying almost parallel to stem; lobes obtuse to subacute and never mucronate at apex, subequal in size, lateral slightly smaller than median; postical leaf bases unarmed; underleaves absent or minute.................................... 3. Barbilophozia barbata

7.  Lateral leaves distinctly narrowed to base, not symmetric, not lying parallel to stem; lobes often mucronate at apex, unequal in size; postical leaf bases with 1-several cilia; underleaves present.

8. Plants often red pigmented (purple or carmine) at shoot apices, leaf bases, and underleaves; central laminal cells 30--40 /um wide; oil-bodies up to 26--28 per cell; underleaves small and indistinct; Greenland……….. 12. Barbilophozia rubescens

8. Plants lacking red pigments; central laminal cells 16--28 /um wide; oil-bodies 2--10 per cell; underleaves large and distinct; widespread.

9. Plants up to 4--4.5(--5) mm wide; leaves usually +/- undulate; apices of leaf lobes mostly with spinose mucros;  postical leaf bases often composed of long and often numerous cilia of strongly elongated cells; gemmae usually absent ……………….................. 10. Barbilophozia lycopodioides

9..Plants (0.5--)1.5--2(--3.3) mm wide; leaves not or slightly undulate; apices of leaf lobes usually acute to mucronate; postical leaf bases composed of few cilia of moderately elongated cells; gemmae usually present, abundant. 7. Barbilophozia hatcheri

6.  Lateral leaves mostly (2--)3-lobed.

10. Underleaves absent or minute, well under 1/5 leaf length.

11. Attenuate, erect gemmiferous shoots with closely appressed leaves with reduced lobes usually present, filiform; laminal cells with usually small and concave-sided trigones; gemmae mostly pale green, in exposed sites partly reddish or vinaceous. ...................................... 2. Barbilophozia attenuata

11. Attenuate, erect gemmiferous shoots absent or present and not filiform; laminal cells often with large, convex-sided trigones. Gemmae sporadically present, red or purplish red.

12. Plants erect or suberect usually growing among mosses, rarely on rock; underleaves absent; mature leaves invariably 3-lobed with sinuses 1/4--1/2 the leaf length.   ……… 4. Barbilophozia bisteadii

12. Plants procumbent and slightly ascending to suberect, usually growing on rocks, occasionally on soil and among mosses; underleaves absent or small and filiform to oblong and bi-lobed; mature leaves 2--3(--4)-lobed, with sinuses 0.25--0.4 the leaf length. .1. Barbilophozia atlantica

10. Underleaves large, up to ½ leaf length or longer.

13. Underleaves usually about or less than half the leaf length. ……………………. 9. Barbilophozia kunzeana (in part)

13. Underleaves more than half the leaf length.

14.  Plants usually large to robust, (1--)1.8--3(--5) wide; sinuses 1/8--2/5 leaf length; gemmae absent.  ………………………………….. 6. Barbilophozia floerkei

14. Plants medium-sized, 0.8--1.1 mm wide; sinuses 1/3--1/2 leaf length; gemmae present or absent …… 8. Barbilophozia hyperborea (in part)

 

 

1. Barbilophozia atlantica (Kaalaas) Müller Frib , Leberm. Eur., 639. 1954

Jungermannia atlantica Kaalaas, Skr. Vidensk.-Selsk. Christiania, Math.-Naturvidensk. Kl. 1898(9): 11. 1898; Orthocaulis atlanticus (Kaalaas) H. Buch

 

Plants 1--3(--)5 cm x 0.4--0.8(--2) mm, prostrate to ascending, rarely erect, usually in mats, sometimes in cushions, green, yellow-green to dark gray-green, or yellow-brown or brownish red, sometimes with a purple tinge. Stems with shoots sometimes weakly attenuate and gemmiferous. Lateral leaves weakly obliquely inserted, erecto-patent or patent, not secund, sub-orbicular to obtrapezoidal, usually complicate-concave, occasionally more or less horizontal and partly convex, not or slightly undulate, 0.6--1.2(--1.5) x 0.4--0.8(--1) mm, unequally (2--)3(--4)-lobed, apices acute to sub-acute, rarely rounded, margins often irregularly slightly sinuous or erose on older leaves, rarely apiculate distally, postical base unarmed or with a small tooth, slime papilla, or short-celled cilium; sinuses 1/8--2/5 leaf length, usually gibbous; central laminal cells 4--5 angular, 20--23 x 28--30 \um, walls thin, trigones large and convex, sometimes concave-sided, oil bodies (3--)5--7(--10) per cell. Underleaves absent or small and filiform to oblong and bi-lobed. Specialized asexual reproduction as sub-orbicular or angular gemmae usually present on non-modified or weakly, erect attenuate shoots, gemmae red or purplish red, 18--28 x 20--42(--56) \um, 1--2-celled. Sexual condition dioicous. Gynoecia more frequent, bracts 2--5 lobed, bracteole 2--3 lobed. Perianth mouth dentate or dentate-ciliate; mature perianths unknown.

 

Exposed to shaded, often calcareous rock, often in crevices, occasionally on soil and among mosses; arctic-alpine; 0--1000 m; Greenland; Nfld. and Labrador (Labr.), Nun., Ont., Que; Maine, N.H; Eurasia.

 

Traits characteristic of Barbilophozia atlantica are the arctic-alpine range, sometimes weakly attenuate gemmiferous shoots, red to purple gemmae, and large leaf cells usually with bulging trigones.

 

2. Barbilophozia attenuata (Martius) Loeske, Verh. Bot. Vereins Prov. Brandenburg 49: 37. 1907

Jungermannia attenuata fo. attenuata Martius, Fl. Crypt. Erlang., 177. 1817; Neoorthocaulis atenuata (Martius) L. Söderström, De Roo & Hedderson; Orthocaulis attenuata (Martius) A. Evans

 

Plants 1--3 (--5) cm x 0.7--1.8 mm, prostrate to sub-erect, in mats, green or yellow-green, sometimes brownish or with a red tinge. Stems commonly developing filiform, innovations from shoot apices. Lateral leaves somewhat obliquely inserted, spreading, sub-quadrate or obtrapezoidal, 0.7--0.9 x 0.7--0.8(--1) mm, (2--)3(--4)-lobed, lobes concave or straight-sided, apices acute to sub-acute, sometimes apiculate, postical base unarmed, sinuses 1/3--1/4 leaf length, rarely weakly gibbous; central laminal cells 4--5 angular, 15--20 x 15--25 \um, walls thin or slightly thickened, trigones absent or small, sometimes large, usually concave, oil bodies (3--)6--10 per cell. Underleaves absent or represented by small cilium. Specialized asexual reproduction as polymorphous gemmae on filiform, often erect innovations bearing imbricate, weakly tri-lobate leaves, gemmae green, sometimes orange or weakly vinaceous in exposed sites; 18--22 x 22--35 \um, 1--2-celled. Sexual condition dioicous. Gynoecia with bracts 3--5 lobed, bracteole 2--4 lobed. Perianth cylindrical or oblong-cylindrical, plicate, mouth laciniate and ciliate.

 

Boulders and ledges, talus and rocky slopes, decaying wood; 0--2000 m; Greenland; Alta., B.C., N.B., Nfld. and Labr. (Nfld., Labr.), N.S., Nun., Ont., Que.; Alaska, Colo., Conn., Idaho, Maine, Mass., Mich., Minn., Mont., N.H., N.Y., N.C., Penn., Tenn., Vt., Va., Wash., W.Va., Wis., Wyo.; Eurasia; Atlantic Islands (Azores).

 

Barbilophozia attenuata commonly has attenuate gemmiferous shoots as does B. atlantica but the attenuation is abrupt and narrow, not gradual and indistinct as in the latter species. Additional distinctive traits are the relative lack of postical cilia and the small laminal cells of B. attenuata.

 

3. Barbilophozia barbata (Schmidel ex Schreber) Loeske, Verh. Bot. Vereins Prov. Brandenburg 49: 37. 1907  F

Jungermannia barbata Schmidel ex Schreber, Spic. Fl. Lips., 107. 1771

 

Plants 3--8 cm x 2--5 mm, prostrate, sometimes ascending distally, in loose mats, dark green or green, sometimes brownish green, rarely brown. Stems without distinctive innovations. Lateral leaves obliquely inserted, wide-spreading, obcuneate-quadrate and nearly symmetric, usually lying almost parallel to stem, only slightly narrowed to the base, up to (0.8--)1--1.8(--1) mm, (3--)4-lobed, lobes +/-triangular, sub-equal in size with lateral ones slightly smaller, slightly concave, or straight-sided, apices obtuse to sub-acute, not mucronate, postical base unarmed, sinuses 1/6--1/4 leaf length, not or slightly gibbous; central laminal cells rounded-polygonal, 20--25 x 23--30 \um, walls thin, trigones absent or small and concave, oil bodies 4--9(--12) per cell. Underleaves absent or minute, subulate or lanceolate, usually 2-fid. Specialized asexual reproduction rare as reddish-yellow, angulate gemmae, 1--2-celled. Sexual condition dioicous. Gynoecia with bracts 3--5 lobed, bracteole 2--4 lobed. Perianth oblong-cylindrical, deeply plicate, mouth crenulate or dentate. Capsule wall 4-stratose.

 

Cliffs and outcrops, crevices and ledges, loamy or rocky banks, humus, peaty depressions; 0--1000 m; Greenland; Miquelon; Alta., B.C., N.B., Nfld. and Labr. (Nfld.), N.S., N.W.T., Nun., Ont., Que., Sask., Yukon; Alaska, Ariz., Colo., Conn., Idaho, Ind., Maine, Mass., Mich., Minn., Mont., N.H., N.M., N.Y., N.C., Penn., Vt., Va., Wash., Wis., Wyo.; Eurasia; Atlantic Islands (Iceland).

 

The nearly quadrate leaves lying almost parallel to stem make Barbilophozia barbata the most easily recognized species in the genus. The triangular leaf lobes lack terminal mucros, and the underleaves are absent or very small.

 

4. Barbilophozia binsteadii (Kaalaas) Loeske, Hedwigia 49: 13. 1909

Jungermannia binsteadii Kaalaas, Skr. Vidensk.-Selsk. Christiana, Math.-Naturvidensk. Kl. 1898(9): 9. 1898; Lophozia binsteadii (Kaalaas) A. Evans; Neoorthocaulis binsteadii (Kaalaas) L. Söderström, De Roo & Hedderson

 

Plants 1--3 cm x 0.5--1.2 mm, erect or suberect usually among mosses, often crowded and more or less caespitose, golden-brown to chestnut brown. Stems without distinctive innovations, seldom with gemmiferous shoots. Lateral leaves obliquely inserted, sub-erect and concave (cup-like), sub-quadrate or narrowly obtrapezoidal, usually imbricate and dorsally more or less secund, 0.6--0.9(--1.1) x 0.7--0.95 mm, (2--)3-lobed, lobes sub-equal in shape, apices acute, postical base unarmed, sinuses 1/4--1/2 leaf length, usually weakly gibbous; central laminal cells rounded-polygonal, 17--23 x (18--)24--30 \um, walls thin or slightly thickened, trigones large, convex, oil bodies mostly 3--6 per cell. Underleaves absent or reduced to slime papillae. Specialized asexual reproduction rare, as vinaceous red or deep metallic purple, angulate gemmae, 1--2-celled. Sexual condition dioicous. Gynoecia with bracts similar to lateral leaves but 3--5 lobed, bracteole large and 2-lobed. Perianth cylindrical-clavate, deeply plicate, mouth with small, ciliate-dentate lobes.

 

Tundra, bogs, peaty depressions, rarely on shaded rock, strongly associated with Sphagnum spp. and Calypogeia sphagnicola; arctic-alpine; 0--2000 m; Greenland; Alta., B.C., N.B., Nfld. and Labr. (Labr.), N.S., Nun., Que., N.W.T., Yukon; Alaska; n Eurasia.

 

Barbilophozia binsteadii is distinguished by its mostly erect plants usually among mosses, the usually 3-lobed, rather cupped leaves with narrow, deep sinuses and narrow lobes, and leaf cells with large trigones. Underleaves are absent.

 

5. Barbilophozia cavifolia (H. Buch & S. Arnell) Stotler & Crandall-Stotler, Bryologist 80: 426. 1977

Orthocaulis cavifolius H. Buch & S. W. Arnell, Memoranda Soc. Fauna Fl. Fen. 26: 71. 1951;

Anastrophyllum cavifolium (H. Buch & S.W. Arnell) Lammes; Lophozia cavifolia (H. Buch & S. W. Arnell) R. M. Schuster

 

Plants 1--3 cm x 0.7--1.1 mm, erect or suberect among mosses, brown to reddish-brown, green in shaded sites. Stems without distinctive innovations. Lateral leaves almost transversely inserted, widely spreading, more or less hemispheric, usually remote to contiguous and antically more or less secund, strongly concave, 0.65--0.9 x 0.5--0.95(--1.2) mm, 2(--3 rarely 4) lobed, lobes sub-equal in shape, apices obtuse, incurved, postical base unarmed, sinuses 1/3--1/4 leaf length, not gibbous; central laminal cells mostly 4-sided and isodiametric or short-rectangular, 22--28 x 25--35(--40) \um, walls thin or slightly thickened, trigones large, usually convex, oil bodies (2--)6--8(--12) per cell. Underleaves usually absent or minute, linear-subulate or linear-lanceolate, usually 2-lobed. Specialized asexual reproduction sparse as reddish brown, angulate gemmae, (1--)2-celled. Sexual condition dioicous. Gynoecia unknown.

 

Tundra, peatlands, amongst other mosses; Arctic; 0--200 m; Greenland; Eurasia.

 

Barbilophozia cavifolia is characterized by mostly 2-lobed leaves that are strongly concave and nearly hemispheric, with laminal cells usually having large, convex-sided trigones. This species resembles robust Sphenolobus minutus from which it is distinguished by (4--5)6--8(--12) oil bodies per cell.

 

6. Barbilophozia floerkei (Weber & D. Mohr) Loeske, Verh. Bot. Vereins Prov. Brandenberg 49: 37. 1907

Jungermannia floerkei F. Weber & D. Mohr. Bot. Taschenb., 410. 1807; Orthocaulis floerkei (F. Weber & D. Mohr) H. Buch

 

Plants 2--5(--10) cm x (1--)1.8--3(--5) mm, prostrate, suberect, or erect, in mats, often intertwined with mosses, pale or yellow- green to dark green, sometimes brownish green. Stems without distinctive innovations. Lateral leaves obliquely inserted, patent, spreading, or sub-squarrose, subquadrate to sub-rectangular, nearly orbicular, or obtrapezoidal, imbricate, concave or convex, sometimes undulate, (0.5--)0.9--2 x (0.5--)1--1.5 mm, (2--)3(--4)-lobed, lobes sub-equal in size, apices obtuse to broadly-acute, postical base with 1--3(--6) relatively short cilia composed of +/-isodiametric cells, sinuses 1/8--2/5 leaf length, weakly to strongly gibbous; central laminal cells more or less rounded-polygonal, 16--20 x 20--26 \um, walls thin, trigones large and slightly convex to minute, oil bodies 2--6(--7) per cell. Underleaves large, 2-fid and divided to near base, short-ciliate near base, lobes lanceolate-acuminate and usually ending in a short cilium. Specialized asexual reproduction absent. Sexual Condition dioicous. Gynoecia with bracts 3--4(--7) lobed, bracteole 2-lobed. Perianth ovate-cylindrical, deeply plicate, mouth ciliolate or dentate.

 

On humus, boulders; 0--2000(--3100) m; Alta., B.C., Nfld. and Labr. (Nfld.), N.S., N.W.T., Nun., Que., Yukon; Alaska, Colo., Maine, Mont., N.H., N.Mex.. Vt., Wash., Yukon; South America (Peru); n. Europe.

 

Barbilophozia floerkei usually has subquadrate to sub-rectangular leaves with short cilia on postical margins, composed of near isodiametric cells, and the underleaves are relatively large. It lacks gemmae.

 

7. Barbilophozia hatcheri (A. Evans) Loeske, Verh. Bot. Vereins Prov. Brandenburg 49: 37. 1907

Jungermannia hatcheri A. Evans, Bull. Torrey Bot. Club 25: 417. 1898

 

Plants 2--5 cm x 1--2(--3) mm, prostrate, suberect, or erect, in patches or mats, green or brown, sometimes brownish green. Stems without distinctive innovations. Lateral leaves obliquely inserted, patent or spreading, shape variable, mostly obtrapezoidal, approximate or imbricate, more or less flat, 1--1.5(--2) x 1--1.3 mm, (3--)4-lobed, lobes sub-equal in size with lateral ones smaller, apices acute or obtuse, usually with distal apiculus, rarely with short mucros, postical base with 2--3(--6) relatively short cilia usually composed of elongate-rectangular cells, sinuses 1/5--2/5 leaf length, gibbous or not; central laminal cells quadrate or hexagonal, 20--26 x 18--25 \um, walls thin, trigones small to medium-sized, slightly convex, oil bodies 2--5(--10) per cell. Underleaves large, usually 2-fid and divided to near base, or lanceolate, 10--16 cells wide proximally, long-ciliate at base, cilia often contorted, lobes lanceolate-acuminate, usually ending in a long cilium. Specialized asexual reproduction usually abundant as reddish brown, angulate gemmae, 2-celled. Sexual condition dioicous. Gynoecia with bracts 3--5-lobed, bracteole 2 or 3-lobed. Perianth ovoid-cylindrical, plicate, mouth short-dentate or short-ciliolate.

 

On cliffs. ledges, acidic or somewhat basic rock, particularly under spruce and fir; 300--3000 m elv.; Greenland; Alta., B.C., Nfld. and Labr. (Nfld. and Labr.), N.S., Nun., Ont., Que., Yukon; Alaska, Calif., Colo., Idaho, Maine, Mich., Minn., Mont., N.H., N.Y., N.C., Oreg., Tenn., Utah, Wash., Wyo.; South America (s Argentina); Atlantic Islands (Falklands); Eurasia; Antarctica.

 

Barbilophozia hatcheri is bipolar in distribution and the only species of the genus to occur in Antarctica. The leaves are usually 4-lobed and obliquely inserted. The leaf lobes are usually apiculate, and the postical bases have short cilia composed of elongate cells. The underleaves are large and bear cilia. This species is often confused with smaller plants of B. lycopodioides. It is also similar to B. rubescens. See comparative notes under those species.

 

8. Barbilophozia hyperborea (R. M. Schuster) Stotler & Crandall-Stotler ex Potemkin, Novosti Sist. Nizsh. Rast. 28: 148. 1992

Lophozia floerkei var. hyperborea R. M. Schuster, Bull. Natl. Mus. Canada 164: 21. 1959; L. hyperborea (R. M. Schuster) R. M. Schuster; L. hyperborea ssp. helophila R. M. Schuster; Neoorthocaulis hyperboreus (R. M. Schuster) L. Söderström, De Roos & Hedderson; Orthocaulis hyperboreus (R. M. Schuster) Konstantinova, Novosti Sist. Nizsh. Rast. 30: 112. 1995

 

Plants 1--2 cm x 0.8--1.1 mm, patent or suberect among other bryophytes, brown to brownish-black, sometimes with a red cast. Stems without distinctive innovations. Lateral leaves obliquely inserted, erect or sub-erect, oblong or obtrapezoidal, patent and imbricate, concave to concave-folded, 0.4--1 x 0.4--0.8 mm, (2--)3(rarely 4)-lobed, lobes sub-equal in size, often longer than wide, incurved, apices usually acute, sometimes obtuse, sometimes with distal apiculus, postical base with 1--3 short teeth, less often with short cilia, sinuses 1/3--1/2 leaf length, weakly to strongly gibbous; central laminal cells usually rounded-polygonal, 20--26 x 20--28(--30) \um, walls thin, with coarse cuticles, trigones small to medium-sized, usually concave, oil bodies (4--)7--15 per cell. Underleaves large, rarely small to minute, 2-fid and divided to near base, toothed, sometimes ciliate or laciniate at base, lobes lanceolate-acuminate, usually ending in a long cilium. Specialized asexual reproduction absent or present. Sexual condition dioicous. Gynoecia with bracts 2--3-lobed, bracteole 2-lobed. Perianth ovoid, weakly plicate, mouth ciliate-dentate and lobulate-cililate.

 

Soil, rock; low elevations; Greenland; Nun.; Alaska.

 

Barbilophozia hyperborea is distinguished in part by brown coloration, leaves oblong or obtrapezoidal, with 1--3 short teeth on postical bases, and laminal cells with a coarse cuticle and many oil bodies per cell, and large underleaves.

 

1. Plants without a red coloration, underleaves large, gemmae absent ................................             ...................................................................................................................................              8a. Barbilophozia hyperborea subsp. hyperborea

1. Plants with a red coloration, underleaves small to minute, gemmae present ...................             ...................................................................................................................................             8b. Barbilophozia hyperborea subsp. helophila

 

8a. Barbilophozia hyperborea subsp. hyperborea

 

Plants without a red coloration. Underleaves usually large. Specialized asexual reproduction absent.

 

Wet sites, soil and shale below snow banks and fen-ice cliffs; moderate elevations; Greenland; Nun. (Ellesmere I.); Alaska; n Eurasia.

 

Specimens of Barbilophozia hyperborea fo. paradoxa R. M. Schuster, known from northern Greenland, apparently differ in having only 2--7 oil bodies per cell, coarse trigones, and nearly smooth laminal cell cuticle.

 

8b. Barbilophozia hyperborea subsp. helophila (R. M. Schuster & Damsholt) R. H. Zander, Phytoneuron 2020-17: 1. 2020.   E

 

Lophozia hyperborea ssp. helophila R. M. Schuster & Damsholt, Phytologia 63: 325. 1987; Neoorthocaulis hyperboreus subsp. helophilus (R. M. Schuster & Damsholt) Stotler & Crandall-Stotler

 

Plants with a red coloration. Underleaves small to minute. Specialized asexual reproduction by gemmae.

 

Wet sites, soil and rock; low to moderate elevations; s Greenland.

 

Barbilophozia hyperborea subsp. helophila is restricted to southern Greenland, and was distinguished by R. M. Schuster and K. Damsholt (1987) from the typical subspecies by the key traits above.

 

SELECTED REFERENCE  Zander, R.H. 2020. A new combination in Barbilophozia for Lophozia hyperborea subsp. helophila (Scapaniaceae, Marchantiophyta) of Greenland, with macroevolutionary justification. Phytoneuron 2020-17: 1–3.

 

9. Barbilophozia kunzeana (Hübener) Müll. Frib., Mitteil. Badischen Landesver. Naturk. Naturschutz, N. F. 4: 431. 1944

Jungermannia kunzeana Hueberner, Hepaticol. Germ., 115. 1834; Lophozia kunzeana (Huebener) A. Evans; Orthocaulis kunzeanus (Huebener) H. Buch; Schljakovia kunzeana (Huebener) Konstantinova & Vilnet

 

Plants 2--5(--8) cm x 0.6--1.5 mm, erect or ascending, solitary or in patches, often intertwined with mosses, yellow-brown to chestnut brown, sometimes bright green. Stems without distinctive innovations. Lateral leaves obliquely to nearly transversely inserted especially distally, patent or spreading, sometimes upper portions nearly erect, sub-quadrate to nearly orbicular, approximate to sub-imbricate, concave or convex, 0.6--0.8(--1) x 0.6--0.9(1.2) mm, 2(-- rarely 3)-lobed, lobes sub-equal in size, apices obtuse to rounded, occasionally acute, rarely apiculate, usually incurved, postical base usually unarmed, rarely with a short cilium or tooth, sinuses (1/6--)1/4 – 1/2 leaf length, gibbous; central laminal cells more or less isodiametric, less often short-rectangular, rounded-polygonal, 17--20 x 21--26 \um, walls thin, trigones small, concave, rarely convex, oil bodies (1--)2--4(--5) per cell. Underleaves small, 2-fid and divided to near base, sometimes undivided, lobes lanceolate, sometimes with 1(--3) basal teeth. Specialized asexual reproduction uncommon, yellow-green, brown, or orange-brown angulate gemmae, 1--2-celled. Sexual condition dioicous. Gynoecia with bracts larger than lateral leaves, 3--4(--6) lobed, bracteole 2-lobed. Perianth oblong-ovoid, plicate, mouth dentate-serrate, teeth ciliate.

 

On wet peat over rock, soil, arctic-alpine, heath; 0--1500 m elev.; Greenland; B.C., N.W.T., Nun., Ont., .Sask., Que., Yukon; Alaska, Colo., Maine, Mich., Minn., N.H., N.Y.; Eurasia.

 

Females plants of Barbilophozia kunzeana are usually larger than male plants. The leaves lack postical cilia and are largely 2-lobed.

 

10. Barbilophozia lycopodioides (Wallroth) Loeske, Verh. Bot. Vereins Prov. Brandenburg 49: 37. 1907

Jungermannia lycopodioides Wallroth, Fl. Crypt. Germ. 1: 76. 1831

 

Plants 2--5(--10) cm x 1.5--3(--5) mm, prostrate, sometimes suberect, in mats, often intertwined with mosses, green, pale green, or stramineous yellow, rarely brownish. Stems without distinctive innovations. Lateral leaves obliquely inserted, spreading, obtrapezoid, asymmetrical, imbricate, usually undulate, sometimes crispate, 2--3 x 1.5--2 mm, (rarely 3--)4-lobed, lobes sub-equal in size, apices obtuse, sometimes broadly acute, usually with 1--3 celled mucros, often longest on postical lobe, postical base with 4--7 long and slender cilia composed of elongate cells, sinuses 1/10--1/4 leaf length, weakly to strongly gibbous; central laminal cells rounded-polygonal or short-rectangular, 20 x 25--28 \um, walls thin, trigones large and slightly convex to minute, oil bodies 2--5(--10) per cell. Underleaves large, 2-fid and divided to near base, rarely lanceolate, strongly ciliate, cilia (3--)5--7(--8) long and tortuous, lobes lanceolate-acuminate and ending in a long cilium. Specialized asexual reproduction rare as reddish-brown, angulate gemmae, 2-celled. Sexual Condition dioicous, usually sterile. Gynoecia with bracts 4--5 lobed, bracteole 2(--4)-lobed. Perianth ovate-cylindrical, plicate, mouth dentate. Capsule wall 3-stratose.

 

On well-drained humus, rock walls, ledges; 0--1500 m elv., Greenland; Alta., B.C., Nfld. & Labr. (Labr.), N.W.T., N.S., Que., Yukon; Alaska, Colo., Mont., N.Mex., Utah. Wash., Wyo.; Eurasia.

 

Barbilophozia lycopodioides has been confused with larger plants of B. hatcheri. The leaves of B. lycopodioides, however, are more shallowly lobed and the lobes broader and more strongly gibbous and are often tipped with a long mucro. Also, the antical leaf margins of B. lycopodioides across the mid-line of the stem, whereas those of B. hatcheri do not extend that far.

 

11. Barbilophozia quadriloba (Lindberg) Loeske, Hedwigia 49: 13. 1909

Jungermannia quadriloba Lindberg in Lindberg & Arnell, Kongl.l Svenska Vetensk.-Akad. Handl. (n. ser.) 23: 55. 1889; Lophozia quadriloba (Lindberg) A. Evans; Orthocaulis quadriloba (Lindberg) A. Evans

 

Plants 0.6--2.5(--4) cm x 0.5--1.8 mm, erect, sometimes sub-erect or procumbent, in tufts or dense patches, fuscous or olive-green or brownish, sometimes with inky cast. Stems without distinctive innovations. Lateral leaves subtransversely inserted, spreading with lobes sub-erect or erect, obtrapezoidal, distant to contiguous, imbricate distally on stems, often concave and cupped, 1--1.5 x 0.5--1.3 mm, often wider than long, (rarely 2--3--)4(rarely--5)-lobed, lobes ovate-triangular, sub-equal in size, sometimes dorsal lobe larger, apices acute, postical base with 1--3 cilia or teeth, antical base less ornamented, with 1--3 teeth or short cilia, or not ornamented, sinuses 2/5--3/5 leaf length, often rounded and open near base on larger leaves, unevenly (wavy) gibbous; central laminal cells rounded-polygonal, 22--25(--35) x 18--24 \um, walls thin or slightly thickened, coarsely papillose with large, low papillae, trigones minute to large, rarely convex, oil bodies 4--9 per cell. Underleaves large, 2-fid and divided to near base, sometimes lanceolate and small, usually long- and short-ciliate at base, lobes linear-lanceolate, acuminate. Specialized asexual reproduction absent. Sexual Condition dioicous, usually sterile. Gynoecia with bracts 4--5 lobed, bracteole 2-lobed. Perianth oblong-obovoid, plicate, mouth lobulate-dentate.

 

The unique trait of large, low papillae on the laminal cell wall cuticles, except in highly reduced forms, is characteristic of Barbilophozia quadriloba. This species is also distinctive in the usually 4-lobed leaves with deep sinuses, although some smaller plants have only 2- or 3-lobed leaves, and gemmae are absent. The var. glareosa, a reduced variant with mostly 3-lobed leaves with lobes little reflexed, was recently reduced to Barbilophozia quadriloba fo. glareosa (E. Jørgensen) Potemkin and is not recognized in the Flora. Likewise, the var. glareosa fo. cephalozielloides R. M. Schuster, an extremely reduced form of the variety, characterized by its 2-lobed lateral leaves, is not recognized. It resembles some species of Cephaloziella but is distinguished by its larger leaf cells and longer and lanceolate-lobed underleaves. Slender forms of Chandonanthus setiformis strongly resemble B. quadriloba but it is a warmer brown and clearly shiny when dry, and has dentate, more deeply divided leaf lobes.

 

On soil, humus; low to moderate elevations; Greenland; North America (Canada, n U.S.A.); Eurasia; Atlantic Islands.

 

1. Central leaf cells averaging 22--25 x 18--19 \um, with small, less distinct, rarely slightly convex trigones .................................. 11a. Barbilophozia quadriloba var. quadriloba

1. Central leaf cells averaging 25--35 x 20--24 \um, with large and distinct, often convex trigones .....................................  11b. Barbilophozia quadriloba var. collenchymatica

 

11a. Barbilophozia quadriloba (Lindberg) Loeske var. quadriloba

Barbilophozia quadriloba var. glareosa (E. Jørgensen) Lammes in T. Koponen, Isoviita &

Lammes

 

Laminal cells averaging 22--25 x 18--19 \um, with small, less distinct, rarely slightly convex trigones.

 

On slopes below snow banks, thin soil or humus over basaltic or calcareous rock; 0--1300 m elv.; Greenland; B.C., Nfld. and Labr. (Nfld.); Nun., Ont.; Alaska, Mich.; Eurasia; Atlantic Islands (Iceland).

 

11b. Barbilophozia quadriloba var. collenchymatica (R. M. Schuster) Stotler & Crand.-Statl. Bryologist 80: 410. 1977

Lophozia quadriloba (Lindb.) A. Evans var. collenchymatica R. M. Schuster, Hepat. Anthocerotae N. Amer. 2: 81. 1969; Schljakovianthus quadrilobus var. collenchymaticus Konstantinova & Vilnet, Arctoa 18: 66. 2009 [2010]

 

Laminal cells averaging 25--35 x 20--24 \um, with large and distinct, often convex trigones.

 

On soil, humus; low to moderate elevations, Greenland.

 

12. Barbilophozia rubescens (R. M. Schuster & Damsholt) Karttunen & L. Söderström, Ann. Bot. Fenn. 29: 120. 1992

Lophozia rubescens R. M. Schuster & Damsholt, Phytologia 63: 325. 1987; Barbilophozia hatcheri var. grandiretis H. Buch ex Lammes

 

Plants 2--4 cm x 2.5--3.5 mm, prostrate or ascending, in loose tufts and patches, yellowish green to light brown, shoot apices, leaf bases, and underleaves usually red pigmented (purple or carmine). Stems without distinctive innovations. Lateral leaves obliquely inserted, densely imbricate with lobes incurved, obtrapezoidal to reniform, concave, 1--1.5 x 1--1.3(--2) mm, 3(--4)-lobed, lobes triangular, sub-equal in size with lateral ones smaller,  apices acute, sometimes obtuse, usually with distal mucro, sometimes short-ciliate, postical base with 4--6(--7) long, contorted thread-like cilia usually composed of elongate-rectangular cells, sinuses 1/5--1/3(--1/2) leaf length, gibbous or not; central laminal cells quadrate to short-rectangular, (30--)32--35(--38) x 35--42/um, walls thin, trigones weakly developed to scarcely bulging, oil bodies 5--28 per cell. Underleaves indistinct, 2-lobed and divided up to ca. 0.9 of length, 4--6 cells wide proximally, with cilia on margins near apex or almost throughout, lobes lanceolate, ending a long cilium. Specialized asexual reproduction by gemmae, vinaceous, angular, 1--2-celled, rare. Sexual condition dioicous. Gynoecia with bracts 2-lobed, bracteole 2-lobed. Perianth ovoid, plicate, mouth with scattered 1--3-celled teeth.

 

On wet acidic or calcareous soil; low elevations; s Greenland; n Europe.

 

As implied by its epithet, Barbilophozia rubescens is distinguished in part by the commonly intensely red shoot apices, leaf bases, and underleaves. The leaves are 3(--4)-lobed, underleaves are present but small, and gemmae few or absent. Barbilophozia rubescens is similar to densely leaved plants of B. hatcheri. The leaves, however, are more shallowly lobed and the lobes broader and more strongly gibbous.

 

 

EXCLUDED

Barbilopholia sudetica (Nees ex Huebener) L. Söderström, De Roo & Hedderson is in FNA treated as Lophozia sudetica (Nees ex Huebener) Grolle.

 

 

ScapaniaceaeBarbilophoziaBarbata_Art_12-1