BFNA Title: Lophozia
Author: V. Bakalin
Date: August 23, 2011
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. LOPHOZIA (Dumortier) Dumortier, Recuel Observ. Jungerm., 17. 1835 [Greek lophos, crest, spike, alluding to angular ends of lobes]

Vadim Bakalin

Jungermannia sect. Lophozia Dumortier, Syll. Jungerm., 52. 1831

Plants ca. 3--50 x 0.4--3 mm, prostrate to ascending, from pale green to rusty brown and purplish. Stems 0.1--0.5 mm in diameter, sparsely branching, transverse section with microcellous layer to  1/2\x stem thickness or not differentiated; rhizoids sparse to common, in indistinct fascicles, colorless to pale brown. Leaves succubous, inserted at various angles, distant to imbricate, spreading to appressed, flat to concave, divided by variously shaped sinus into 2(--3--4) lobes; lobes entire; cells commonly collenchymatous, walls usually thin, but sometimes thickened; oil bodies 2--40 per cell, biconcentric or not; underleaves commonly absent. Specialized asexual reproduction by gemmae in masses at apices of leaf-lobes or shots, absent in some species. Sexual condition mainly dioicous, rarely autoicous or paroicous. Androecia intercalary, 1--4-androus, antheridial stalk 1--2-seriate. Gynoecia terminal, female bracts in 1 pair. Perianth exerted 1/2--3/4\x its length, mainly cylindric, complicated at mouth, mouth dentate to ciliate and lobulate; perigynium totally absent. Capsule oval, walls 3--5-stratose, exterior wall with nodular thickenings, interior wall with annular thickenings, reddish brown. Elaters 2-spiraled, reddish brown. Spores spheric, 7--18 \um, papillose, dark brownish.

Species ca. 60 (23 in the flora): North America, Central America, South America, Eurasia, Africa, Australia, Oceania, Antarctica.

SELECTED REFERENCES  Bakalin V.A. 2005. Monograficheskaya obrabotka roda Lophozia (Dumort.) Dumort. s. str. Moscow.  Hong W. S. 2002. The distribution of Lophozia in North America west of the hundredth meridian. Lindbergia. 27: 49--62.  Schuster R. M. 1969. Lophozia (Dumort.) Dumort. In: R. M. Schuster. 1969. Hepaticae and Anthocerotae of North America. New York and London. Vol. 2.

1. Stem cross section with microcellous layer near ventral side.

2. Leaf lobes obtuse to rounded, cells of midleaf with many (to 50) small oil-bodies, underleaves filiform, commonly present but early deciduous, rarely absent, capsule wall (4--)5-stratose . . . XXe. Lophozia subg. Obtusifolium, p. XX (in part)

2. Leaf lobes acute, or leaves only slightly retuse, cells of midleaf with (1--)3--40 oil bodies per cell, underleaves mainly absent, if present then simple, lanceolate, nearly the same width as stem, not deciduous, capsule wall 3(--4)-stratose.

3. Cells of midleaf usually 25--35 \um wide; gemmae colorless to red and purplish, rarely brown (if brown, then plants whitish, gemmae with long persistent, single, commonly biconcentric oil-body), perianth complicated at mouth, dentate to ciliate along margin . . . XXa. Lophozia subg. Lophozia, p. XX

3. Cells of midleaf usually 18--22 \um wide; gemmae brown; perianth narrowed to the mouth, mouth dentate, commonly ornamented by sharply elongated cells . . . XXc.Lophozia subg Sudeticae, p. XX

1. Stem cross section lacking a microcellous layer near ventral side.

4. Leaves subtransversely inserted, imbricate, stem only 0.10--0.15 mm in diameter, underleaves always absent . . . XXb. Lophozia subg. Isopaches, p. XX

4. Leaves inserted at various angles, not imbricate, stem as a rule more than 0.2 mm in diameter, underleaves present or absent.

5. Leaves 2--3(--4)-lobed, obliquely inserted, underleaves more or less regular in shape, plants monoicous, gemmae absent . . . XXd. Lophozia subg. Protolophozia, p. XX

5. Leaves 2--3-lobed, obliquely to subhorizontally inserted, underleaves commonly irregular, plants dioicous, gemmae present, sometimes absent, but in that case leaves nearly horizontally inserted and 2-lobate.

6. Leaves almost horizontally inserted, 2-lobed, lobes obtuse to rounded . . . XXe. Lophozia subg. Obtusifolium, p. XX (in part)

6. Leaves obliquely to subtransversely inserted, 2--3(--4)-lobed, acute . . . XXc. Lophozia subg. Sudeticae, p. XX

XXa. LOPHOZIA (Dumortier) Dumortier subg. LOPHOZIA

Plants ca. 4--40 x 0.6--3 mm, ascending to prostrate, pale green-whitish to yellowish green, brown and purplish brown. Stems 0.1--0.5 mm in diameter, sparsely branching, transverse section with microcellous layer to 1/2\x stem thickness, frequently present, but sometimes nearly absent depending on growth habit or ecological condition, outer cells of stem in cross section not or slightly thickened. Leaves transversely to subhorizontally inserted, more or less distant, spreading, 2(3--4)-lobed; cuticle smooth to papillose; underleaves rarely present, simple, lanceolate, nearly the same length as leaves. Specialized asexual reproduction by angular gemmae, with slightly to sharply thickened angles, colorless to red, occurring in all species. Sexual condition dioicous, rarely monoicous or paroicous. Capsule oval, wall 3(--4)-stratose, Spores spheric, 9--18 \um.

Species 22 (17 in the flora): North America, South America, Eurasia, Africa, Australia, Oceania, Antarctica.

1. Stems in cross section with thickened superficial cell walls along margin; mature gemmae colorless to greenish; antheridial bracts with entire margins . . . XXa.1. Lophozia sect. Lophozia, p. XX

1. Stems in cross section with thin cell walls throughout; mature gemmae red, reddish, carrot-red, brown, brown-orange to purplish; antheridial bracts sometimes with additional antical teeth . . . XXa.2. Lophozia sect. Excisae, p. XX

XXa.1. Lophozia sect. Lophozia

Lophozia sect. Guttulatae Schljakov; Lophozia sect. Heteromorpha R. M. Schuster; Lophozia sect. Savicziae Bakalin

Plants ca. 4--40 x 0.6--2 mm, ascending to prostrate, pale green-whitish to yellowish-green, brown and purplish-brown. Stems with outer cells in cross section slightly thickened. Leaves transversely to subhorizontally inserted, more or less distant, spreading, 2(--4)-lobed, oil-bodies biconcentric or not; cuticle smooth to papillose. Specialized asexual reproduction by colorless to greenish gemmae [brownish]. Sexual condition dioicous [monoicous]. Antheridial bracts with entire margins.

Species ca. 14 (10 in the flora): North America, Eurasia, Antarctica.

1. Leaves commonly 2--3-lobed, crispate-undulate; gemmae colorless, but sometimes violet. . . 3. Lophozia heteromorpha

1. Leaves uniformly 2-lobed, only very rarely 3-lobed, flattened to concave but margin neither undulate nor crispate; gemmae colorless to greenish, rarely with rose or brownish tinge.

2. Leaves only shallowly retuse at apex, base distinctly decurrent dorsally; perianth fusiform . . . 4. Lophozia pacifica

2. Leaves clearly lobed with acute to weakly acute lobes, base not decurrent dorsally, perianth cylindric to subclavate.

3. Plants whitish, ascending; leaves erect-spreading with lobes deflexed from shoot apex; ventral side of stem brownish or the same color as the dorsal; epixylous . . . 1. Lophozia ascendens

3. Plants greenish to brown and purplish, prostrate to erect in dense patches; leaf lobes erect or rarely deflexed; ventral side of stem brown to purplish; epixylous or not.

4. Plants generally bright-green, more rarely yellowish, particularly when epixylous; mature leaves longer than wide.

5. Leaves flattened, leaf insertion line nearly straight, leaf lobes near apex deflexed. . . 7. Lophozia silvicola

5. Leaves grooved, leaf insertion line incurved from very oblique on ventral side to subtransverse dorsally, leaf lobes near apex upturned.

6. Midleaf cells ca. 22--26 \um wide with bulging trigones, near margin sometimes confluent; cuticle commonly striolate . . . 2. Lophozia guttulata

6. Midleaf cells ca. 22--33 \um wide, trigones triangular to convex, or if cell width less than 26 \um then trigones concave; cuticle commonly smooth.

7.  Midleaf cells commonly more than 26 \um wide, trigones triangular to convex, oil-bodies biconcentric; perianth with teeth more than (3--)4 cells in length . . . 8. Lophozia silvicoloides

7. Midleaf cells commonly less than 26 \um wide, trigones concave, oil-bodies non-biconcentric; perianth with 1--2(--5)-celled teeth . . . 9. Lophozia ventricosa

4. Plants generally yellow-brownish to brown, purplish and rusty-red; mature leaves wider than long.

8. Leaves grooved, leaf base semi-sheathing and leaf insertion line incurved from very oblique ventrally to subtransverse dorsally; leaf lobes near apex turned up to form ob-pyriform cap…. 9. Lophozia ventricosa

8. Leaves flattened or cape-shaped but not grooved and base not semi-sheathing base, leaf insertion line nearly straight, leaf lobes near apex slightly incurved inside, leaves not forming ob-pyriform cap.

9. Leaves without trace of red or purple pigmentation, even dorsally.

10. Leaves obliquely inserted, frequently with deeply brownish colored leaf lobes near apex; oil-bodies biconcentric ….        6. Lophozia schusterana

10. Leaves subtransversely inserted, pigmentation of leaf lobes not different; oil-bodies non-biconcentric….     10. Lophozia wenzelii

9. Leaves commonly with red, purple or purplish pigmentation, at least dorsally.

11. Midleaf cells mostly wider than 29 \um; gemmae penta- to polygonal with protruding angles; oil-bodies commonly more than 20 per cell, at least 50% of them biconcentric ….           5. Lophozia savicziae

11. Midleaf cells mostly narrower that 29 \um; gemmae 3--4(--5)-angular with protruding angles; oil-bodies commonly less than 20 per cell, biconcentric….          7. Lophozia silvicola

Key to living perianthous plants

1. Oil-bodies biconcentric.

2. Perianth mouth laciniate or ciliate, with cilia to (3--)4--6 cells in length….          8. Lophozia silvicoloides

2. Perianth mouth dentate to shortly laciniate, with cilia not more than 3(--4) cells in length.

3. Oil-bodies more than 20 per cell, smooth, some homogenous non-biconcentric (not more than 50%); leaves subtransversely inserted….           5. Lophozia savicziae

3. Oil-bodies less than 20 per cell, granulate; leaves obliquely to subhorizontally inserted.

4. Leaves mostly flattened and longer than wide; Arctic to Boreal zones….                                                                 7. Lophozia silvicola

4. Leaves concave, wider than long; chiefly Arctic …       6. Lophozia schusterana

1. Oil-bodies non-biconcentric.

5. Perianth fusiform; leaves retuse, not distinctly 2-lobed….       4. Lophozia pacifica

5. Perianth cylindric to subclavate; leaves always distinctly lobed.

6. Perianth mouth laciniate to ciliate-laciniate with cilia more than (4--)5 cells in length….                                                                    1. Lophozia ascendens

6. Perianth mouth dentate to ciliate with cilia shorter than 4 cells in length.

7. Leaves polymorphous 2--3(--4)-lobed, lanceolate, underleaves occasionally present; gemmae and distal portion of perianth sometimes with violet tint….                       3. Lophozia heteromorpha

7. Leaves uniformly 2(--3)-lobed; underleaves absent; no violet tint in gemmae or distal portion of perianth.

8. Leaves subtransversely inserted and oriented, strongly concave to cupped, wider than long; shoots not red pigmented….                                          10. Lophozia wenzelii

8. Leaves obliquely inserted and oriented (rarely subtransversely, but then longer than wide), flattened to grooved, rarely concave, wider than long to longer than wide; shoots often reddish pigmented.

9. Leaves longer than wide, characteristically grooved, trigones strongly convex to bulging; leaf cuticle sometimes striolate; perianth mouth ciliate with cilia 3--4 cells length ….                          2. Lophozia guttulata

9. Leaves wider than long (occasionally longer than wide, but then trigones concave and leaves flattened), flattened to concave and grooved, trigones concave to convex; leaf cuticle smooth; perianth mouth dentate or ciliate with cilia 3(--5) cells in length….      9. Lophozia ventricosa

1. Lophozia ascendens (Warnstorf) R. M. Schuster, Bryologist, 55: 180. 1952

Sphenolobus ascendens Warnstorf, Hedwigia 57: 63. 1915; Lophozia ciliata Damsholt, Soederstroem & Weibull

Plants ca. 4--10 x 0.6--1 mm, ascending, very rarely prostrate, pale green-whitish and yellowish white. Stems 0.15--0.35 mm in diameter, sparsely branching, transverse section with microcellous layer to  1/4\x stem thickness, sometimes absent in attenuate tips; rhizoids sparse to common, in indistinct fascicles, colorless to pale brownish. Leaves transversely inserted, distant, spreading, erect-spreading at the ends of lobes, deflexed away from the stem, flat, rectangular to ovate, 0.5--0.7 x 0.3--0.6 mm, 2-lobed, equal lobes divided by U-shaped to crescentic sinus descending to 1/4--2/5\x leaf length; lobes entire, angular to horn-like; cells of midleaf rectangular to polygonal, 23--28 x 20--25 \um, in base to  40 \um; cuticle smooth, walls thin, trigones small, triangulate to slightly convex; oil bodies 4--10(--14) per cell, spheric to ovoid, 3--5 x 4--8 \um, finely granular, grayish, non-biconcentric; underleaves rarely present, lanceolate, nearly the same length with leaves. Specialized asexual reproduction by gemmae in masses at apices of leaf-lobes, greenish to colorless, commonly triangular to rectangular in outline, (12--)15--22 x 11--17 \um, 2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, often with gemmae at tips, male bracts 2 pairs, similar to leaves but inflated in the base, 1-androus, antheridial stalk 1-seriate. Gynoecia terminal, female bracts in 1 pair, rectangular to trapezoidal, sometimes dentate, ca. 0.6 x 0.35--0.7 mm, divided to 1/3\x length, connate; bracteole present, lanceolate, sometimes shortly 2-lobed to 1/3\x length, ca. 0.5 x 0.2--0.3 mm, connate with one of bracts to 2/3\x length. Perianth exerted to 3/4\x length, cylindric to slightly rhomboidal, complicated at mouth, mouth ciliate to lobulate with lobules and cilia to  7--8 cells in length, generally 1-stratose. Seta 5 mm, capsule oval, wall 3-stratose, exterior wall with nodular thickenings, interior wall with annular thickenings, reddish brown. Spores spheric, 9--11 \um.

Decaying wood, predominantly wet dark coniferous forests (Picea, Abies, etc.), sometimes bark of decaying trees; 0--3200 m; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., N.S., Ont., Que., Yukon; Alaska, Colo., Idaho, Maine, Mich., Minn., Mont., N.H., N.Y., Oreg., S.Dak., Vt., Wash., Wis., Wyo.; n Eurasia.

2. Lophozia guttulata (Lindberg & Arnell) Evans, Proc. Washington Acad. Sci. 2: 302. 1900

Jungermannia guttulata Lindberg & Arnell, Musci As.-Bor. 51. 1889

Plants ca. 3--10 x 0.8--1 mm, prostrate, yellowish pale green to brownish light green. Stems 0.1--0.4 mm in diameter, sparsely branching, transverse section with microcellous layer to 1/5--1/4\x stem thickness, ventral surface brown-purple to purple-brown-blackish; rhizoids sparse to common, in indistinct fascicles, colorless to purplish near origin area. Leaves obliquely inserted, insertion line incurved from nearly horizontal on ventral side to nearly transverse on the dorsal, distant to imbricate, spreading, turn up at the ends of lobes, slightly lengthways sulcate, rectangular, 0.7--1 x 0.5--0.6 mm, 2-lobed, subequal lobes divided by angular to gibbous sinus descending to 1/5--1/4\x leaf length; lobes entire, triangular to arching; cells of midleaf 4--5-angular, 25--27 x 22--25 \um, in base to 25--30(--35) x 25 \um; cuticle smooth to papillose, walls thin, trigones bulging, sometimes confluent; oil bodies 4--10(--12) per cell, spheric to ovoid, 4--5 x 5--10 \um, finely granular, grayish, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of shoots, sometimes absent, greenish to colorless, commonly triangular to rectangular, (12--)15--22 x 11--17 \um, 2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts in 2--3 pairs, similar to leaves but inflated basally, 1(--2)-androus, antheridial stalk 1-seriate. Gynoecia terminal, female bracts in 1 pair, obovate to rectangular, entire, 1.2--1.3 x 0.8--1 mm, divided to 1/3\x length, connate; bracteole present, lanceolate to narrowly rectangular and shortly 2-lobed to 1/3\x length, connate with one of bracts to 1/3\x length. Perianth exerted to 2/3\x length, cylindric to somewhat clavate, complicated at mouth, mouth dentate to ciliate with cilia to 4(--8) cells in length, generally 1-stratose. Seta ca. 5 mm, capsule oval, walls 3-stratose, exterior walls with nodular thickenings, interior walls with annular thickenings, reddish brown. Spores 11--13 \um in diameter.

Decaying wood, mainly dark coniferous forests, wet peaty soil or peat-moss in boggy areas, more rarely humus soil and fine-grained ground along lakes in forested areas; 0--3300 m; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Ont., Que., Yukon; Alaska, Calif., Colo., Conn., Idaho, Iowa, Maine, Mass., Mich., Minn., Mont., N.H., N.Mex., N.Y., Oreg., Utah, Vt., Wash., Wis., Wyo.; Central America; n Eurasia.

3. Lophozia heteromorpha R. M. Schuster & Damsholt, Schuster, Hep. Anth. North Amer. 2: 507. 1969

Plants ca. 5--25 x 1--2 mm, prostrate, pale to bright green or greenish brown and blackish, frequently with violet-colored tips. Stems 0.3--0.8 mm in diameter, sparsely branching, transverse section with microcellous layer to 1/3\x stem thickness or absent, ventral surface brown to black-brown and purple-brown; rhizoids sparse to common, in indistinct fascicles, colorless to pale violet near origin area. Leaves obliquely inserted, distant to contiguous, spreading, undulate, polymorphous from orbicular to subquadrate and trapezoidal, 0.8--1 x 0.8--1.4 mm, 2--3(--4)-lobed, subequal lobes divided by crescentic to gibbous sinus descending to 1/5--2/5\x leaf length; lobes entire, triangular to arching; cells of midleaf 4--5-angular, 18--30(--35) x 17--27 \um; cuticle smooth, walls thin to somewhat thickened, trigones distinct, triangular to concave, oil bodies unknown; underleaves sometimes present, filiform to lanceolate to  200--300 \um. Specialized asexual reproduction rare, by gemmae at apices of shoots, colorless to pale violet, commonly triangular to rectangular in outline, 14--19 x 14--24 \um, (1--)2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia unknown. Gynoecia terminal, female bracts in 1 pair, obovate to rectangular, entire, 1.2--1.3 x 0.8-1 mm, divided to 1/3\x length into 2--3 lobes, connate, bracteole present, lanceolate to narrowly rectangular and shortly 2-lobed to 1/3\x length, connate with one of bracts to 2/3\x length. Perianth exerted to 2/3\x length, cylindric to somewhat ob-clavate, complicated at mouth, mouth dentate with few teeth 2(--3) cells in length, generally 1-stratose. Other features unknown.

Among mosses and liverworts in dense patches, as well as in disturbed areas, data on ecology very incomplete; elevation unknown; Greenland; Alaska; n  Eurasia

4. Lophozia pacifica Bakalin, Bryologist 114: 302, fig. 1. 2011

Plants 10--20 x 2--2.3 mm, prostrate to ascending, bright green to yellowish green without traces of reddish coloration except ventral side of stem, dorsiventrally flattened. Stems 0.35--0.4 mm in diameter, sparsely branching, transverse section with microcellous layer to 1/5\x stem thickness or absent, ventral surface red-brown; rhizoids sparse to nearly absent, originated in dorsal side of stem at right angle. Leaves obliquely to nearly horizontally inserted, clearly decurrent on dorsal side, obliquely spreading, somewhat concave with lobes incurved inwardly, distant, oval to widely ovate, 1.2--1.5 x 1.3--1.4 mm, shallowly retuse; cells of midleaf 4--5-angular, 28--35 x 25--28 \um, basally to 50 \um; cuticle smooth but at base distinctly verrucose, walls thin, trigones convex to bulging; oil-bodies 4--8 per cell, finely papillose, irregularly ellipsoidal, 4--5.5 x 3--4.7 mm; underleaves absent. Specialized asexual reproduction by solitary gemmae at apices of shoot, greenish to colorless, commonly 4--5-angular in outline, 15--25 x 15--20 \um, 2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 2--3 pairs, trapezoidal, inflated in the base, 1--2-androus, 90--120 \um, antheridial stalk 1-seriate. Gynoecia terminal with one or two innovations, female bracts in 1 pair, obovate, entire, 1.3--1.5 x 1.1-1.3 mm, divided 1/8\x length into 2 lobes by crescentic sinus, connate, bracteole present, lanceolate ca. 1.3 x 0.35-0.45 mm, connate with one of bracts to 2/3\x length. Perianth exerted to 5/6\x length, fusiform, constricted to mouth, ca. 3 x 1 mm, mouth dentate with sharply elongated 1-celled teeth, generally 1-stratose. Sporophyte unknown.

Ecology incompletely known, apparently confined to tundra belt of low mountains, wet mineral to humus soil, partial shade to full sun; 1500--1800 m; Wash.

5. Lophozia savicziae Schljakov, Novosti Sist. Nizsh. Rast. 10: 299. 1973

Lophozia silvicola var. grandiretis H. Buch & S.W. Arnell; L. ventricosa var. grandiretis (H. Buch & S.W. Arnell) R. M. Schuster & Damsholt

Plants ca. 5--20 x 1.5--2 mm, usually prostrate, rarely ascending, yellow-brown to rusty brown. Stems (0.2--)0.25--0.3 mm in diameter, sometimes slightly transversely oval in cross section, sparsely branching, transverse section with microcellous layer to  2/3\x stem thickness, red-brown in color, ventral surface of stem brown-purple to purple-brown, or purple-black; rhizoids sparse to common, in indistinct fascicles, colorless to purplish near origin area. Leaves subtransversely inserted, insertion line more or less straight, distant to imbricate, spreading, turned up at the ends of lobes, strongly concave to cup-shaped, orbicular to trapezoidal, 0.8--1.1 x 1.1--1.5 mm, sometimes 2-stratose in base to 1/10\x length, 2-lobed, unequal lobes divided by crescentic to angular or gibbous sinus descending to 1/5\x leaf length; lobes entire, triangular to arching; cells variable in size, in midleaf 5--6-angular, 25--45 x 25--40 \um, but mostly exceeding 30 \um wide; cuticle smooth to papillose, walls thin, trigones usually bulging, sometimes confluent; oil bodies 15--40 per cell, spheric to ovoid, ca. 2--5 \um, finely granular to nearly smooth, light bluish, more than 60% of oil-bodies are biconcentric; underleaves very rarely present, lanceolate. Specialized asexual reproduction by gemmae in masses at apices of shoots, sometimes absent, greenish to colorless, commonly 5-angular, more rarely 4-angular to polygonal in outline, (18--)22--30 x 15--30(--33) \um, 1--2-celled, with strongly thickened and protruding angles. Sexual condition dioicous. Androecia unknown. Gynoecia terminal, female bracts in 1 pair, obovate to rectangular, entire, 1.2--1.5 x 1.5--2 mm, divided to 1/3\x length into 2--3 lobes, rarely connate; bracteole narrowly rectangular, shortly 2-lobed to 1/4\x length, 0.8--0.9 x 1.1--1.2 mm, connate with one of bracts to 1/3\x length. Perianth exerted to 5/6\x length, cylindric to somewhat subclavate, complicated at mouth, mouth dentate to shortly lobulate with teeth to 2(--3) cells in length, generally 1-stratose. Sporophyte unknown.

Northern tundra, sometimes in northern mountains, among mosses or in pure mats in wet habitats, mainly places with percolated water, often somewhat alkaline rocks; 0--1000 m; Greenland; Alaska; n Eurasia.

6. Lophozia schusterana Schljakov, Novosti Sist. Nizsh. Rast. 12: 320. 1975

Plants ca. 5--30 x 1--2 mm, prostrate to erect, in dense patches, yellowish to pale green and peculiarly brownish at the tips, commonly laterally compressed. Stems 0.3--0.48 mm in diameter, sparsely branching, transverse section with microcellous layer to  1/3\x stem thickness, ventral surface brownish purple to reddish brown; rhizoids sparse to rarely common, in indistinct fascicles, colorless to purplish near origin area. Leaves obliquely to horizontally inserted, distant to imbricate, commonly appressed, rarely spreading, rectangular to nearly orbicular, antically secund, 0.9--1.1 x 0.9--1.2 mm, 2-lobed, subequal lobes divided by crescentic to angular sinus descending to 1/5--1/4\x leaf length; lobes entire, triangular to arching; cells of midleaf 4--5-angular, 24--32 x 23--26 \um; cuticle smooth, walls thin, trigones convex to bulging; oil bodies 8--18(--30) per cell, spheric to ovoid, 4--6 x 5--10 \um, finely granular, grayish, biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of shoots, sometimes absent, greenish to colorless, commonly triangular to rectangular in outline, (18--)21--25 x 18--20 \um, 1--2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 2--3 pairs, similar to leaves but inflated in the base, 1(--2)-androus, antheridial stalk 1--2-seriate. Gynoecia terminal, female bracts in 1 pair, obovate to rectangular, entire, 1.2--1.3 x 0.8--1 mm, divided to 1/3\x length into 2--4 lobes, connate; bracteole present, lanceolate, connate with one of bracts to 1/3\x length, 1--1.5 x 0.5 mm. Perianth exerted to 2/3\x length, cylindric to somewhat clavate, complicated at mouth, mouth dentate with remote 1--2-celled teeth, generally 1-stratose. Sporophyte unknown.

Restricted to tundra zone, not exceeding mountain tundra belt in boreal (taiga) areas, overhanging rock in snow-bed communities, crevices, among mosses and other liverworts, rarely submerged in temporary streams or on moist soil on steep slopes; 0--2200 m; Greenland; B.C., Nfld. and Labr. (Labr.), Nunavut; Alaska, Mich.; n Eurasia.

Lophozia schusterana is L. groenlandica in the sense of Schuster (1969) but not Jungermannia groenlandica Nees.

7. Lophozia silvicola H. Buch, Rep. 18. Scand. Naturalist Congr., 228. 1929

Lophozia ventricosa var. silvicola (H. Buch) Jones

Plants ca. 10--20 x 1.7--2(--2.2) mm, prostrate to creeping, yellowish green to bright green. Stems 0.25--0.3 mm in diameter, sparsely branching, transverse section with microcellous layer to 1/3--2/3\x stem thickness, ventral surface brown to purple-brown; rhizoids sparse to common, in indistinct fascicles, colorless to yellowish and brownish. Leaves commonly very obliquely inserted but rarely subtransverse, insertion line nearly straight, distant, spreading, flattened, deflexed from the apex, rectangular, 1--1.5 x 0.7--1 mm, 2-lobed, subequal lobes divided by crescentic to angular sinus descending to 1/5--1/4\x leaf length; lobes entire, acute, triangular to almost horn-like, sometimes ending by 2 superposed cells; cells of midleaf 5--6-angular, 28--30 x 25--30 \um, in base to 50 \um; cuticle smooth, walls thin, trigones concave, triangular to rarely bulging; oil bodies 7--18(--20) per cell, spheric to ovoid, 4--7 x 5--10 \um, finely granular, grayish, biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at lobe apices of distal leaf, greenish to colorless, commonly 3--5-angular in outline, 15--18 x 12--17 \um, 2-celled, with thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 2--4 pairs, similar to leaves but inflated in the base, 1--2-androus, antheridial stalk 2-seriate. Gynoecia terminal, innovations common, female bracts in 1 pair, obovate to rectangular, entire, 1.5--2(--2.2) x 1.3--1.7 mm, divided to 1/3\x length into 2--3 lobes, connate, bracteole present, lanceolate, connate with one of bracts to 1/3\x length. Perianth exerted to 2/3\x length, cylindric, complicated at mouth, mouth dentate with 2--3-celled teeth, generally 1-stratose. Seta ca. 10--15 mm, capsule oval, wall 3--4-stratose, exterior wall with nodular thickenings, interior wall with annular thickenings, reddish brown. Elaters ca. 60 x 5--6 \um. Spores spheric, 10--13 \um, papillose, dark brown.

Dark coniferous forest communities on decaying wood and in moss mats on forest floor, wet places, also tundra and boggy areas over moss mats, humus and rarely pure peat, steep slopes with disturbed vegetation cover; 0--2900 m; Greenland; Alta., B.C., Nfld. and Labr., N.W.T., N.S., Ont., Que.; Alaska, Calif., Colo., Idaho, Maine, Mass., Mich., Minn., N.H., N.Y., N.C., Tenn., Vt., Va., Wash., Wis., Wyo.; n Eurasia.

8. Lophozia silvicoloides N. Kitagawa, J. Hattori Bot. Lab. 28: 276. 1965

Plants ca. 5--30 x 0.5--2(--2.5) mm, prostrate with ascending tips, pale to bright green sometimes brownish. Stems 0.25--0.4 mm in diameter, sparsely branching, transverse section with microcellous layer to  1/3--2/3\x stem thickness, ventral surface red-brown to purple-black; rhizoids sparse to common, in indistinct fascicles, colorless to reddish near origin area. Leaves commonly obliquely inserted, rarely subtransverse, insertion line curved from nearly horizontal on dorsal surface to subtransverse on the ventral, distant, spreading, flattened, deflexed from the apex, rectangular to ovoid, 0.5--1.3 x 0.4--1.2 mm, 2-lobed, subequal (dorsally secund) lobes divided by crescentic to angular and U-shaped sinus descending to 1/4--1/3(--2/5)\x leaf length; lobes entire, acute to somewhat acute, triangular to arching; cells of midleaf 5--6-angular, (22--)28--40 x 25--45 \um, in base to  50 \um; cuticle smooth, walls thin to slightly thickened, trigones triangular to bulging; oil bodies 10--20(--25) per cell, spheric to ovoid, 3--7 x 3--8 \um, finely granular, light bluish, biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at lobe apices of distal leaf, greenish to colorless, commonly 3--5-angular in outline, 15--28 x 13--27 \um, 2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 3--5 pairs, similar to leaves but inflated in the base, 1--2-androus, antheridial stalk 2-seriate. Gynoecia terminal, innovations common, female bracts in 1 pair, obovate to rectangular, entire, 1.5--2(--2.2) x 1.3--1.7 mm, divided to 1/3\x length into 2--4 lobes, connate, bracteole present, lanceolate, connate with one of bracts to 1/3\x length. Perianth exerted to 4/5\x length, cylindric, to subclavate, 3--4 x 1--1.2 mm, complicated at mouth, mouth dentate to ciliate and lobulate with lobules 2--3 cell in base and 5--7 cells in length, generally 1-stratose. Spores 13--18 \um in diameter, papillose, dark brown.

Arctic-montane species with suboceanic distribution, tundra, over mosses in dense patches in mesic areas; 0--100 m; Alaska; n Eurasia.

9. Lophozia ventricosa (Dickson) Dumortier, Recuel Observ. Jungerm., 17. 1835

Jungermannia ventricosa Dickson, Pl. Crypt. Brit., fasc. 2: 14. 1790; Lophozia subapiculata R. M. Schuster & Damsholt

Plants ca. 10--20 x 1--2(--2.2) mm, prostrate to ascending and nearly erect in dense patches, bright green to yellowish green, yellowish brown, and reddish brown. Stems 0.13--0.3 mm in diameter, sometimes slightly transversely oval, sparsely branching, transverse section with microcellous layer to  1/3--1/4\x stem thickness, ventral surface brown to purple-brown; rhizoids sparse to common, in indistinct fascicles, colorless to yellowish and brownish near origin. Leaves commonly obliquely inserted but rarely subtransverse, insertion line curved from nearly horizontal on dorsal surface to subtransverse on the ventral, distant, spreading, concave to flattened, turned up the apex, rectangular, obtrapezioidal to suborbicular, 0.8--1.2 x 0.75--1.2 mm, usually 2-lobed, unequal or subequal lobes divided by angular to gibbous sinus descending to 1/5--1/4\x leaf length; lobes entire, acute, triangular to arching, sometimes ending by 2 superposed cells; cells of midleaf 5--7-angular, 25--31 x 23--28 \um, in base to  45 \um; cuticle smooth, rarely verrucose, walls thin, trigones concave, triangular to bulging; oil bodies 7--18(--20) per cell, spheric to ovoid, 4--7 x 5--10 \um, finely granular, grayish, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at apex of shoots, greenish to colorless, commonly 3--4(--5)-angular in outline, 15--25 x 12--20 \um, (1--)2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 2--3 pairs, similar to leaves but inflated in the base, 1--2-androus, antheridial stalk 1--2-seriate. Gynoecia terminal, female bracts in 1 pair, obovate to rectangular and obtrapezioidal, entire, 1--2 x 1--2 mm, divided to 1/3\x length into 2--3 lobes, connate; bracteole present, lanceolate, connate with one of bracts to 1/3\x length. Perianth exerted to 3/4\x length, cylindric, complicated at mouth, mouth dentate with 2--4(--5)-celled teeth, generally 1-stratose. Seta ca. 10--15 mm, capsule oval, wall 3--4-stratose, exterior wall with nodular thickenings, interior wall with annular thickenings, reddish brown. Elaters  ca. 100--175 x 6--8 \um. Spores spheric, 10--13 \um, papillose, dark brown to reddish brown.

Varieties 2 (2 in the flora): North America, Eurasia, Antarctica.

1. Leaves longer than wide, divided by angular to U-shaped sinus, trigones in midleaf cells concave . . . 9a.Lophozia ventricosa var. ventricosa

1. Leaves wider than long, divided by gibbous sinus, trigones in midleaf cells convex to bulging . . . 9b.Lophozia ventricosa var. longiflora

9a. Lophozia ventricosa var. ventricosa

Plants ca. 10--20 x 1.8--2 mm, prostrate to ascending and nearly erect in dense patches, bright green to yellowish green, yellowish brown. Stems with ventral surface brown to purple-brown. Leaves spreading, flattened or rarely undulate, turned up the apex, rectangular, obtrapezioidal to orbicular, 0.8--1.2 x 0.75--1.2 mm, usually 2-lobed, unequal or subequal lobes divided by angular sinus descending to 1/5--1/4\x leaf length; lobes entire, acute, triangular, sometimes ending by 2 superposed cells; cuticle smooth, walls thin, trigones concave to triangular.

Among mosses, lichens, dwarf-shrubs, steep slopes in mesic conditions, tundra zone or belt, decaying wood, forest floor, places with disturbed vegetation cover (both native and anthropogenic), also crevices of various rock, banks of streams and lakes; 0--3300; Greenland; Alta., B.C., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Yukon; Alaska, Ariz., Calif., Colo., Conn., Idaho, Maine, Mass., Mich., Minn., Mont., Nev., N.H., N.Y., N.C., Oreg., R.I., Tenn., Utah, Wash., Wis., Wyo.; n Eurasia.

9b. Lophozia ventricosa var. longiflora (Nees) Macoun, Cat. Canad. Pl. 7: 17. 1902

Jungermannia longiflora Nees, Naturg. Eur. Leberm., 2: 95. 1836

Plants ca. 10--20 x 1.8--2.2 mm, prostrate to ascending and nearly erect in dense patches, yellowish green to yellowish brown and reddish brown. Stems ventral surface brown to purple-brown. Leaves spreading, concave to cup-shaped, turned to  the apex, orbicular, 0.8--1.2 x 0.75--1.2 mm, usually 2-lobed, subequal lobes by angular or more usually gibbous sinus descending to 1/5--1/4\x leaf length; lobes entire, acute, triangular to arching, walls thin, trigones convex to bulging.

Common in both zonal and mountain tundra, drying pools, among mosses (mainly Sphagnum), rarely on rocks, also dark coniferous forest on decaying wood, peaty turfs; 0--4000 m; Greenland; B.C., Nfld. and Labr. (Labr.), Ont., Que.; Alaska, Calif., Colo., Idaho, Maine, Minn., Mont., N.H., Wis.; n Eurasia; Central America; Antarctic.

10. Lophozia wenzelii (Nees) Stepheni, Spec. Hepat. 2: 127, fig. 135. 1901

Jungermannia wenzelii Nees, Naturgesch. Eur. Leberm., 2: 6, fig. 58. 1836

Plants ca. 5--40 x 0.2--3 mm, erect in dense patches, small form creeping, yellow-brown, rarely with purplish colored tips of shoots (high Arctic phases). Stems 0.15--0.5 mm in diameter, sparsely branching, transverse section with microcellous layer to  1/3\x stem thickness, ventral surface brownish to black-brown, without reddish pigmentation; rhizoids sparse to rarely common, in indistinct fascicles, colorless to brownish near origin area. Leaves subtransversely inserted, distant to (more frequent) imbricate, commonly appressed, nearly orbicular, sometimes antically secund, (0.2--)0.3--1(--1.2) x (0.2--)0.35--1.3(--1.5 mm, 2-lobed, subequal lobes divided by crescentic to rarely angular sinus descending to 1/5--1/4\x leaf length; lobes entire, triangular to arching; cells of midleaf 4--6-angular, 20--30(--35) x (15--)20--28(30) \um; cuticle smooth, walls thin, trigones concave to convex; oil bodies (2--)4--10(--15) per cell, spheric to ovoid, 4--8 x 5--10 \um, finely granular to almost smooth, grayish, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of shoots, sometimes absent, greenish to colorless, in sunny forms some of them can acquire pale rose or slightly brownish pigmentation, commonly triangular to rectangular in outline, (16--)20--26(--28) x (12--)14--21(--25) \um, 1--2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts to 15 pairs, but commonly only with 5--6 ones, similar to leaves but inflated in the base, 1--2-androus. Gynoecia terminal, female bracts in 1 pair, obovate to transversely rectangular and orbicular, entire, 1.2--1.3 x 1.2--1.5 mm, divided to 1/3\x length into 2--3(--4) lobes, connate; bracteole present, lanceolate to 2-lobate, connate with one of bracts to 2/3\x length, ca. 1 x 0.5 mm. Perianth exerted to 2/3\x length, cylindric, complicated at mouth, mouth dentate with remote 1--2-celled teeth, generally 1-stratose. Spores 10--13 \um in diameter.

Varieties 5 (3 in the flora): North America, Eurasia.

1. Leaves subtransversely to obliquely inserted, divided by angular to gibbous sinus into unequal lobes . . . 10b.Lophozia wenzelii var. groenlandica

1. Leaves subtransversely inserted, divided by angular or crescentic sinus into equal to subequal lobes.

2. Plants erect in dense patches, wider than 1 mm, rich peat moss bogs . . . 10a.Lophozia wenzelii var. wenzelii

2. Plants creeping to ascending, narrower than 0.8 mm, wet soil . . . 10c.Lophozia wenzelii var. lapponica

10a. Lophozia wenzelii var. wenzelii

Plants ca. 5--30 x 1--3 mm, erect in dense patches, yellow-brown. Stems 0.3--0.5 mm in diameter, sparsely branching, transverse section with microcellous layer to 1/3\x stem thickness. Leaves subtransversely inserted, imbricate, commonly appressed, nearly orbicular, equal lobes divided by crescentic sinus descending to 1/5-1/4\x leaf length; lobes entire, obtuse.

Sphagnum bogs, wet mossy tundra, among mosses and other liverworts, 0--3500 m; Greenland; Alta., B.C., Nfld. and Labr., N.W.T., Ont., Que., Sask., Yukon; Alaska, Calif., Colo., Idaho, Maine, Mont., N.H., N.Mex., Oreg., Wash., Wyo.; n Eurasia.

All old and many new records of var. wenzelii could not be checked, but it is apparent that not less than half of them are misidentifications, mainly of L. ventricosa, so the distribution needs additional work.

10b. Lophozia wenzelii var. groenlandica (Nees) Bakalin, Arctoa 10: 213. 2001

Jungermannia groenlandica Nees in Gottsche, Lindenberg & Nees, Syn. Hep., 114. 1844; Lophozia groenlandica (Nees in Gottsche, Lindenberg & Nees) Macoun

Plants ca. 5--40 x 1--3 mm, erect in dense patches or creping on soil, yellow-brown to brown. Stems 0.3--0.5 mm in diameter, sparsely branching, transverse section with microcellous layer to 2/5\x stem thickness. Leaves subtransversely to oblique inserted, imbricate to distant, orbicular or trapezoidal, dorsally secund unequal lobes divided by angular to gibbous sinus descending to 1/5-1/4\x leaf length; lobes entire, triangular.

Dry tops of hummocks in bogs, mesic cliffs, places with disturbed vegetation cover, humus, unforested areas; 0--1800 m; Greenland; B.C.; Alaska, Wash.; n Eurasia.

 10c. Lophozia wenzelii var. lapponica H. Buch & S.W. Arnell, Svensk Bot. Tidskr. 44(1): 81. 1950

Plants ca. 5--30 x 0.2--0.8 mm, creping on soil, yellow-brown to brown and blackish, rarely with purple tips. Stems 0.15--0.3 mm in diameter, sparsely branching, transverse section with microcellous layer to 2/5\x stem thickness. Leaves subtransversely inserted, imbricate, orbicular, subequal lobes divided by angular sinus descending to 1/4\x leaf length; lobes entire, triangular.

Mainly soil in tundra or tundra belt, rarely dark coniferous forest, damp soil or rocks, 0--2700 m, Greenland; N.W.T., Que.; Calif.; n Eurasia.

XXa.2. LOPHOZIA sect. EXCISAE R. M. Schuster, Hep. Anth. North Amer., 2: 511. 1969

Plants ca. 4--50 x 0.6--2 mm, ascending to prostrate, pale green-whitish to yellowish green, brown and purplish brown. Stems with outer cells in cross section thin. Leaves transversely to subhorizontally inserted, more or less distant, spreading, 2-lobed in exceptional cases 3--4-lobed (some leaves on the stem); oil-bodies non-biconcentric; cuticle smooth. Specialized asexual reproduction by gemmae: mature always red to purple, immature sometimes greenish to colorless. Sexual condition dioicous, paroicous, autoicous; antheridial bracts frequently with additional tooth near dorsal side.

Species 8 (7 in the flora): North America, South America, Eurasia, Africa, Australia, Oceania, Antarctica.

1. Plants monoicous or rarely dioicous and if so lax-leaved, soft-textured and bearing (2--)3--4-celled gemmae . . . 11.Lophozia excisa

1. Plants dioicous, not soft-textured, gemmae (1--)2-celled.

2. Plants whitish, small, restricted calcareous sites; oil bodies in leaf cells sometimes biconcentric; gemmae brown with commonly long persistent 1--2 oil bodies per cell . . . 15.Lophozia perssonii

2. Plants green to brownish and brown, not whitish, growing in calcareous sites or not; oil bodies in leaf cells never biconcentric; gemmae red to green, rarely brownish, with many quickly disappear oil-bodies.

3. Plants attenuate with spreading to erect-spreading leaves; ends of lobes deflexed from the apex . . . 12.Lophozia longidens

3. Plants prostrate to erect in dense patches, never ascending; lobes flattened, sometimes crispate-undulate.

4. Gemmae greenish to brownish, concolorous with lobe ends pigmentation; calcareous sites . . . 14.Lophozia pellucida

4. Gemmae greenish (immature only) to deep red and purplish, in color different from that of lobe ends; acidic sites, rarely calcareous.

5. Plants dark brown; gemmae 5--6-polygonal purple; cells of midleaf mostly more than 30 \um wide . . . 17.Lophozia rubrigemma

5. Plants deep-green to brownish, rarely dark brown; gemmae 3--5-angular, rarely 6-angular, but in this case not purple; cells of midleaf mostly less than 30\um wide.

6. Lobes divided by crescentic or U-shaped sinus, gemmae rusty-red . . . 16.Lophozia polaris

6. Lobes divided by angular to gibbous sinus, mature gemmae purple.

7. Oil-bodies granulate . . . 13.Lophozia propagulifera

7. Oil-bodies botryoidal . . . 16b.Lophozia polaris var. sphagnorum

11. Lophozia excisa (Dickson) Dumortier, Recuel Observ. Jungerm., 17. 1835

Jungermannia excisa Dickson, Pl. Crypt. Brit., fasc. 3: 11. 1793.

Plants ca. 10--30 x 1--2.1 mm, prostrate, rarely ascending, yellowish green to yellow-brownish or rarely bright green. Stems 0.15--0.4 mm in diameter, sparsely branching, transverse section with microcellous layer to  1/4--2/3\x stem thickness, sometimes undifferentiated, ventral surface brown to purple-brown; rhizoids dense, in indistinct fascicles, colorless to brownish near origin area. Leaves obliquely inserted, but sometimes line insertion becomes almost transverse on dorsal side, distant to imbricate, spreading to erect-spreading or laterally appressed to the stem, turn up at the ends of lobes, sometimes slightly undulate, rectangular to trapezoidal, 0.55--1 x 0.55--1 mm, 2-lobed, subequal to unequal acute lobes divided by angular to gibbous sinus descending to 1/4--1/3\x leaf length; lobes entire, triangular to arching; cells of midleaf 4--5-angular, 25--32 x 23--30 \um, in base to  ca. 40 x 35 \um; cuticle smooth, walls thin, trigones concave and small, rarely bulging; oil bodies 10--30 per cell, spheric to oval, 3--5 \um in diameter to 3--5 x 4--8 \um if ellipsoidal, granulate, colorless, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae sparse at apices of shoots, frequently absent, greenish when immature to red and purple, commonly 4--5-angular in outline with distinctly protruding angles, 25--38 x 20--28 \um, 2(--3--4)-celled. Sexual condition paroicous. Androecia below gynoecia, male bracts 2--3 pairs, similar to leaves but inflated in the base, sometimes with additional tooth near dorsal base, 1--3(--4)-androus. Gynoecia terminal sometimes with subfloral innovations, female bracts in 1 pair, obovate to rectangular, entire to messy dentate or undulate, 1.2--1.3 x 0.8--1.3 mm, divided to 1/3\x length into 3--4 lobes, connate, bracteole present, lanceolate to narrowly rectangular and shortly 2-lobed to 1/3\x length, connate with one of bracts to 2/3\x length. Perianth exerted to 2/3\x length, cylindric to somewhat pyriform, complicated at mouth, mouth dentate with sparse 1--2-celled teeth, generally 1-stratose. Seta ca. 5 mm, capsule oval, wall 3-stratose, exterior wall with nodular thickenings, interior wall with annular thickenings, reddish brown. Spores 12--20 \um in diameter.

Varieties 3 (2 in the flora): North America, Eurasia, South America, Australia, Antarctica (?).

1. Plants somewhat rigid; stem cross section orbicular or nearly orbicular; specialized asexual reproduction by 2-celled gemmae . . . 11a.Lophozia excisa var. excisa

1. Plants soft-textured; stem cross-section transversely oval; specialized asexual reproduction by 3--4-celled gemmae . . . 11b.Lophozia excisa var. elegans

11a. Lophozia excisa var. excisa

Lophozia excisa var. infuscata R. M. Schuster; L. excisa var. succulenta R. M. Schuster

Plants somewhat rigid. Stems cross-section orbicular or nearly orbicular. Specialized asexual reproduction by 2-celled gemmae.

Tundra, also frequent in mesic conditions in forested areas, exposed and wet soil, among mosses in dense mats, dry humus, snow-bed and snow-fed habitats, steep slopes to river; 0--3300 m; Greenland; Alta., B.C., Man., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Yukon; Alaska, Calif., Colo., Conn., Idaho, Maine, Mich., Minn., Mont., N.H., N.J., N.Y., N.C., Oreg., R.I., Wash., Wis., Wyo.; South America; Eurasia; Australia; Oceania (New Zealand); Antarctica.

11b. Lophozia excisa var. elegans R. M. Schuster, Hep. Anth. North Amer., 2: 522. 1969

Plants soft-textured. Stems cross-section transversely oval. Specialized asexual reproduction by 3--4-celled gemmae.

Northern tundra, mesic conditions among mosses; 0--100 m; Greenland; Nunavut; n Eurasia.

12. Lophozia longidens (Lindberg) Macoun, Geol. Survey Canada: Cat. Canad. Pl. 7: 18. 1902

Jungermannia longidens Lindberg, Bot. Not., 27. 1877

Plants ca. 5--15 x 0.6--1 mm, ascending, rarely prostrate, green to bright green, rarely brownish. Stems 0.17--0.35 mm in diameter, sparsely branching, transverse section with microcellous layer to  1/3--1/2\x stem thickness, sometimes absent in attenuate tips; rhizoids sparse to common, in indistinct fascicles, colorless to pale brownish near origin. Leaves transversely or (rarely, in shady forms) oblique inserted, distant, spreading, erect-spreading at the ends of lobes, deflected away from the stem, flattened, rectangular to ovate, 0.4--0.8 x 0.3--0.55 mm, 2(--3)-lobed, equal lobes divided by U-shaped to semilunate sinus descending to 1/3--1/4\x leaf length; lobes entire, angulate to horn-like; cells of midleaf rectangular to rarely pentagonal, 23--27 x 22--26 \um, in base to 30 \um; cuticle smooth, walls thin, trigones small, concave to triangular; oil bodies 3--12(--17) per cell, spheric, 3--7 \um in diameter to ovoid, 3--5 x 4--9 \um, finely granular, nearly colorless to grayish, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of leaf-lobes, red-brown to brownish and chestnut, rarely greenish in shade forms, commonly 3--4(--5)-angular in outline, 20--30(--40) x 13--24(--36) \um, (1--)2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, often with gemmae at tips, male bracts 3 pairs, similar to leaves but inflated in the base, 1-androus, antheridial stalk 1-seriate. Gynoecia terminal with subfloral innovations, female bracts in 1 pair, rectangular to trapezoidal, sometimes dentate, but commonly entire, ca. 0.6 x 0.35--0.7 mm, divided to 1/3\x length into 2--4 lobes, connate; bracteole present, lanceolate, sometimes shortly 2-lobed to 1/3\x length, ca. 0.5 x 0.2--0.3 mm, connate with one of bracts to 2/3\x length. Perianth exerted to 3/4\x length, ca. 2--2.5 x 0.7 mm, cylindric, complicated at mouth, mouth ciliate to lobulate with lobules and cilia to 5--7 cells in length, generally 1-stratose. Seta 5 mm, capsule oval, walls 3-stratose, exterior walls with nodular thickenings, interior walls with annular thickenings, reddish brown. Elaters ca. 100--120 x 7.5 \um. Spores spheric, 11--13 \um.

Tundra to dark coniferous forest, rarely broad-leaved woods, on rocks of various composition, soil, among mosses, decaying wood, steep slopes to waterways, also places with disturbed vegetation cover; 0--3500 m; Greenland; Alta., B.C., Nfld. and Labr., N.W.T., N.S., Ont., Que., Yukon; Alaska, Colo., Conn., Idaho, Maine, Mass., Mich., Minn., Mont., N.H., N.Y., Oreg., Vt., Wash., Wis., Wyo.; n Eurasia.

 13. Lophozia propagulifera (Gottsche) Stephani, Exped. Antarct. Belge, Bot. 3: 4. 1901.

Jungermannia propagulifera Gottsche, Neumayer, Int. Polarforsch., Deutsch. Exped. 2: 451. 1890; Lophozia latifolia R. M. Schuster

Plants ca. 10--30 x 1.2--1.7 mm, prostrate, but frequently ascending, yellowish brown to yellowish green or rarely bright green. Stems 0.25--0.35 mm in diameter, sparsely branching, slightly transversely oval in cross section, with microcellous layer to 1/2\x stem thickness, cell walls distinctly thickened along margin, ventral surface brown to purple-brown; rhizoids sparse to dense, in indistinct fascicles, grayish. Leaves obliquely to nearly horizontal inserted, appressed to the stem, ends of lobes turned to  apex, sometimes slightly undulate, rectangular to trapezoidal, 0.5--0.7 x 0.5--0.8 mm, 2-lobed, subequal to unequal, acute, lobes divided by angular to gibbous sinus descending to 1/3--2/5\x leaf length; lobes entire, triangular to arching; cells of midleaf 5--6-angular, 28--32 x 23--30(--32) \um, in base to  ca. 43 \um in diameter; cuticle smooth, walls thin to somewhat thick, trigones concave and small; oil bodies 4--25 per cell, spheric 3--6 \um in diameter to oval 3--5 x 4--8 \um, granulate, colorless, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae sparse at apices of shoots, frequently absent, greenish (commonly as admixture, but sometimes to 90%) to purple, commonly 4--5-angular in outline with weakly protruding angles, 25--38 x 20--28 \um, 2-celled. Sexual condition dioicous, autoicous and (rarely) paroicous. Androecia intercalary, male bracts 4(--5) pairs, similar to leaves but inflated in the base, sometimes with additional tooth near dorsal base, 1--2-androus. Gynoecia terminal sometimes with subfloral innovations, female bracts in 1 pair, obovate to rectangular, entire to undulate, 1.2--1.3 x 0.8--1.3 mm, divided to 1/3\x length into 3--4 lobes, connate; bracteole present, lanceolate to narrowly rectangular and shortly 2-lobed to 1/3\x length, connate with one of bracts to 2/3\x length. Perianth exerted to 3/4\x length, cylindric to rhomboidal, complicated at mouth, mouth dentate with sparse 1--2-celled teeth, generally 1-stratose. Seta ca. 5 mm, capsule oval, wall 3-stratose, exterior wall with nodular thickenings, interior wall with annular thickenings, reddish brown. Spores 12--20 \um in diameter.

Damp moss mats, tundra or more rarely forested areas, bogs and rocky cliffs, exposed or partly shaded places; elevation unknown; Greenland; Nfld. and Labr. (Labr.), N.W.T., Nunavut; Alaska, Calif., Minn., Wash.; Eurasia; Antarctica.

 14. Lophozia pellucida R. M. Schuster, Canad. J. Bot. 39: 978. 1961

Lophozia pellucida var. minor R. M. Schuster

Plants ca. 5--30 x 1--2.5 mm, prostrate, yellowish brown to yellowish green or rarely brown. Stems 0.2--0.45 mm in diameter, sparsely branching, slightly transversely oval in cross section, with microcellous layer to  1/4\x stem thickness, commonly weakly developed and indistinct, cell walls distinctly thickened along margin, ventral surface brown; rhizoids sparse to dense, in indistinct fascicles, colorless. Leaves obliquely to nearly horizontal inserted, distant and spreading, rarely appressed to the stem, flattened, ends of lobes peculiarly more deeply colored (commonly distinctly brownish), trapezoidal to transversely oval, 0.5--1.3 x 0.5--1.5 mm, 2-lobed, subequal to unequal acute lobes divided by angular to gibbous sinus descending to 1/3--2/5\x leaf length; lobes entire, triangular to arching; cells of midleaf 4--6-polygonal, 30--45 x 25--40 \um, in base to  ca. 50 \um in diameter; cuticle smooth, walls thin to somewhat thickened, trigones concave and small; oil bodies (5--)9--18(--25) per cell, spheric 4--5.5 \um in diameter to oval 6.5--9 x 4.5--5.5 \um, finely granulate, colorless, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae sparse at apices of shoots and lobe ends, frequently absent, concolorous with leaf lobes, green to brownish or brown, (22--)25--40(--45) x (20--)22--32(--36) \um, 2-celled. Sexual condition dioicous [autoicous and paroicous]. Androecia intercalary, male bracts 3--4 pairs, similar to leaves but inflated in the base, rarely with additional tooth near dorsal base. Gynoecia terminal sometimes with subfloral innovations, female bracts in 1 pair, obovate to rectangular, entire to undulate, 1.2--1.3 x 0.8--1.8 mm, divided to 1/3\x length into 2--4 lobes, connate, bracteole present, lanceolate to narrowly rectangular and shortly 2-lobed to 1/3\x length, connate with one of bracts to 2/3\x length. Perianth exerted to 3/4\x length, cylindric to rhomboidal, complicated at mouth, mouth dentate with sparse 1--2-celled teeth, generally 1-stratose. Sporophyte unknown.

Soil, disturbed vegetation cover, also on mosses in crumbly mats, mainly confined to calcareous sites, rarely on acid silty deposits; elevation unknown; Greenland; Alta., Nunavut; Alaska; n Eurasia.

15. Lophozia perssonii H. Buch & S.W. Arnell in: H Buch, Bot. Not., 381. 1944

Plants ca. 2--10 x 0.5--1 mm, ascending, pale green, brownish to whitish. Stems 0.15--0.3 mm in diameter, sparsely branching, orbicular in cross section, with microcellous layer to  2/5\x stem thickness, cell walls very thin, ventral surface brownish or undifferentiated; rhizoids sparse to dense, in indistinct fascicles, grayish. Leaves subtransversely to slightly oblique inserted, imbricate or spreading, ends of lobes turned to  apex or deflexed away the stem, rectangular to ovate, 0.4--0.7 x 0.3--0.7 mm, 2-lobed, subequal lobes divided by U-shaped or crescentic sinus descending to 1/6--1/3\x leaf length; lobes entire, triangular to nearly horn-like; cells of midleaf 5--6-angular, 21--40 x 20--30 \um; cuticle smooth, walls very thin, trigones concave, very small or absent; oil bodies 3--10 per cell, spheric 3--8 \um in diameter to oval 3--5 x 4--9 \um, granulate, colorless, sometimes biconcentric; underleaves absent. Specialized asexual reproduction by gemmae common at shoot apices, brown to brownish, commonly 3--5-angular in outline with weakly protruding angles, 15--21(--23) x 12--15(--19) \um, 1--2-celled, with long-persistent large single oil-body in each cell. Sexual condition dioicous. Androecia intercalary, male bracts 2--3 pairs, similar to leaves but concave at the base, 1--2-androus. Gynoecia terminal, female bracts in 1 pair, obovate to rectangular, entire to weakly dentate, divided to 1/3\x length into 2 lobes, connate, bracteole present, lanceolate to narrowly rectangular and shortly 2-lobed to 1/3\x length, connate with one of bracts to 2/3\x length. Perianth exerted to 2/3\x length, cylindric to cone-shaped, complicated at mouth, mouth ciliate-dentate with sparse 5--7-celled cilia, generally 1-stratose. Capsule oval, exterior walls with nodular thickenings, interior walls with annular thickenings, reddish brown. Spores 14--18 \um in diameter.

Mesic to xeric limestone cliffs or dry basic clayish deposits; 0--200 m; Greenland; Alaska; n Europe.

16. Lophozia polaris (R. M. Schuster) R. M. Schuster & Damsholt, Meddel. Gronland 199: 131. 1974

Lophozia alpestris ssp. polaris R. M. Schuster, Hep. Anth. North Amer., 2: 614. 1969

Plants ca. 3--30 x 0.6--2.5 mm, prostrate, but frequently ascending in dense patches, green to yellowish brown-green and brown. Stems 0.25--0.5 mm in diameter, sparsely branching, slightly transversely oval in cross section, with microcellous layer to 1/2\x stem thickness, cell walls distinctly thickened along margin, ventral surface brown to purple-brown; rhizoids sparse to nearly absent, in indistinct fascicles, grayish. Leaves obliquely to nearly horizontally inserted, appressed to the stem, ends of lobes upturned, sometimes slightly undulate, rectangular to trapezoidal, 0.4--1.3 x 0.4--1.6 mm, 2-lobed, subequal nearly acute to obtuse lobes divided by crescentic, U-shaped or angular sinus descending to 1/6--1/4(--1/3--2/5)\x leaf length; lobes entire, triangular; cells of midleaf 5--6-angular, (20--)25--35 x (18--)23--35 \um; cuticle smooth, walls thin to somewhat thick, trigones concave and small; oil bodies (3--)6--20 per cell; underleaves absent. Specialized asexual reproduction by gemmae sparse at apices of shoots, rarely on ends of lobes before apex, frequently absent, red, rusty-red to carrot-red, commonly 4--5-angular in outline with weakly protruding angles, 18--32 x 14--28 \um, 1--2-celled. Sexual condition dioicous. Androecia intercalary, male bracts 4--7 pairs, similar to leaves but inflated in the base. Gynoecia terminal without subfloral innovations, female bracts in 1 pair, trapezoidal to transversely oval, entire to slightly undulate, ca. 1.5 x 2 mm, divided to 1/3--2/5\x length into 3--4 lobes with strongly deflexed sinus, connate, bracteole present, 2-lobed to 1/3\x length, connate with one or both bracts to 1/2\x the length. Perianth exerted to 1/2\x its length, cylindric to cone-shaped, complicated at mouth, mouth dentate with sparse 1--2-celled teeth, generally 1-stratose. Sporophytes unknown.

Varieties 2 (2 in the flora): North America, n Eurasia.

1. Lobes divided by crescentic or U-shaped sinus descending to 1/6--1/4\x leaf length; oil-bodies 6--17 per cell, granulate, spheric 3.5--6(--8) \um in diameter to oval (4--)6--8 x 3--6 \um. . . . 16a.Lophozia polaris var. polaris

1. Lobes divided by angular to somewhat gibbous sinus descending to 1/3--2/5\x leaf length; oil-bodies 3--14 per cell, botryoidal, oval 3--5 x 4--6 \um. . . . 16b.Lophozia polaris var. sphagnorum

16a. Lophozia polaris var. polaris

Leaves 2-lobed, subequal nearly acute to obtuse lobes divided by crescentic or U-shaped sinus descending to 1/6-1/4\x leaf length; oil-bodies 6--17 per cell, granulate, spheric 3.5--6(--8) \um in diameter to oval (4--)6--8 x 3--6 \um.

Predominantly basic conditions over mosses and other liverworts in wet places, frequently in percolate water, more rarely cliffs and ledges, banks of sluggishly flowing streams; 0--1000 m; Greenland; Nunavut; Alaska; n Eurasia.

16b. Lophozia polaris var. sphagnorum (R. M. Schuster) R. M. Schuster & Damsholt, Meddel. Gronland 199: 134. 1974

Lophozia alpestris ssp. polaris f. sphagnorum R. M. Schuster, Hep. Anth. North Amer. 2: 619. 1969

Leaves 2-lobed, subequal nearly acute to obtuse lobes divided by angular to somewhat gibbous sinus descending to 1/3--2/5\x leaf length; oil-bodies 3--14 per cell, botryoidal, oval 3--5 x 4--6 \um.

Mostly over Sphagnum, bogs and mossy tundras; 0--500 m; Greenland; Alaska; n Eurasia.

17. Lophozia rubrigemma R. M. Schuster, Hep. Anth. North Amer. 2: 621. 1969

Plants ca. 3--10 x 0.7--1 mm, prostrate, but frequently ascending in dense patches, yellowish brown to brown. Stems 0.25--0.3 mm in diameter, sparsely branching, slightly transversely oval in cross section, microcellous layer to  1/2\x stem thickness, cell walls distinctly thickened along margin, ventral surface brown to purple-brown; rhizoids sparse to nearly absent, in indistinct fascicles, grayish. Leaves obliquely inserted, distant, spreading to appressed to the stem and slightly concave, ends of lobes turned away of stem, sometimes slightly undulate, rectangular to trapezoidal, 0.4--1.2 x 0.4--1.2 mm, 2-lobed, subequal nearly acute to obtuse lobes divided by crescentic, U-shaped or angular sinus descending to 1/3--2/5\x leaf length; lobes entire, triangular; cells of midleaf polygonal to orbicular, (28--)30--38(--43) x 28--36(--45) \um; cuticle smooth, walls somewhat thick, trigones concave and small; oil bodies 10--30 per cell, granulate, spheric 3--5 \um in diameter to shortly oval 3--4.5 x 4--9 \um; underleaves absent. Specialized asexual reproduction by gemmae sparse at apices of shoots, rarely on ends of lobes before apex, red to deep purple-red, commonly (5--)6-polygonal in outline with protruding angles, 25--35 x 25--30 \um, 2-celled. Sexual condition dioicous. Androecia unknown. Gynoecia terminal without subfloral innovations, female bracts in 1 pair, trapezoidal to transversely oval, entire to slightly undulate, ca. 1.5 x 2 mm, divided to 1/4--1/3\x length into 2--3(--4) lobes with strongly deflexed sinus, connate, bracteole present, 2-lobed with sinus descending to 1/3\x length, connate with both of bracts to 1/2\x length. Perianth exerted to 1/2\x its length, cylindric to conic, complicated at mouth, mouth dentate with sparse 2--3-celled teeth, generally 1-stratose. Sporophytes unknown.

Acidophilic, common in Arctic deserts, more rare but also common in some areas of tundra, exposed sites, places with disturbed or light vegetation cover; elevation unknown; Greenland; Nunavut; Alaska; n Eurasia.

XXb. LOPHOZIA subg. ISOPACHES (H. Buch) R. M. Schuster, Amer. Midl. Nat. 45(1): 56. 1951

Isopaches H.Buch, Mem Soc. F. Fl. Fenn., 8: 283. 1933

Plants ca. 3--15 x 0.6--3 mm, prostrate, loosely adhering to substratum, decolorated to deep green, but commonly brown to reddish brown. Stems 0.1--0.15 mm in diameter, sparsely branching, transverse section without microcellous layer, outer cells of stem in cross section clearly thickened, rhizoids conspicuously large. Leaves transversely to obliquely inserted, imbricate, rarely distant, spreading, 2-lobed; cuticle smooth; underleaves rarely present, simple, lanceolate, nearly the same length with leaves. Specialized asexual reproduction by angular gemmae, with sharply thickened angles. Sexual condition paroicous, autoicous to dioicous. Capsule oval, walls 2(--3)-stratose, Spores 12--15 \um in diameter.

Species 4 (3 in the flora): North America, South America, Eurasia, Australia, Oceania (New Zealand).

1. Plants monoicous (occasionally paroicous, very rarely with antheridial subfloral innovations), female bracts entire, perianth mouth lobulate-ciliate, plants strongly aromatic . . . 19. Lophozia bicrenata

1. Plants dioicous, female bracts dentate or entire, perianth mouth crenulate or ciliate-dentate, plants not or slightly aromatic.

2. Female bracts entire, perianth mouth crenulate . . . 20.Lophozia decolorans

2. Female bracts sharply dentate, perianth mouth ciliate-dentate . . . 18. Lophozia alboviridis

18. Lophozia alboviridis R. M. Schuster, Hep. Anth. North Amer. 2: 487. 1969

Plants ca. 3--7 x 0.7--1 mm, prostrate, whitish green to brownish and purplish in upper area. Stems 0.11--0.15 mm in diameter, sparsely branching, orbicular in cross section, with homogenous inner cells, outer cells distinctly thickened, ventral surface brownish; rhizoids dense, in tangled fascicles, colorless to brownish, closely attaching plants to substratum. Leaves subtransversely to obliquely inserted, appressed to the stem, imbricate, ends of lobes turned to  apex, orbicular, 0.5--0.7 x 0.5--0.7 mm, 2-lobed, subequal to unequal acute lobes divided by angular to crescentic sinus descending to 1/4\x leaf length; lobes entire, triangular, acute; cells of midleaf 5--6-angular with rounded angles, 20--24 x 20--28 \um; cuticle smooth, walls clearly thick, trigones concave and small; oil bodies 10--15 per cell, spheric ca. 5--6 \um in diameter to oval 5--6 x 6--8(--10) \um, granulate, colorless, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae common at apices of shoots, reddish yellow to reddish brown, commonly 4--5-angular in outline, with protruding angles, 24--38 x 24--28 \um, 2-celled, red-brown. Sexual condition dioicous. Androecia unknown. Gynoecia terminal, female bracts in 1 pair, obovate to nearly orbicular, sharply dentate, divided to 1/3\x length into 2--3(--4) lobes, connate, bracteole present, lanceolate, connate with one of bracts to 2/3\x length. Perianth mouth lobulate and spinose-dentate with teeth to 2(--3) cells in length, generally 1-stratose. Sporophyte unknown.

Bare, cleaved soil in Arctic polygonal tundra (data incomplete); Greenland; n Eurasia.

19. Lophozia bicrenata (Schmidel ex Hoffmann) Dumortier, Recuel Observ. Jungerm., 17. 1835

Jungermannia bicrenata Schmidel ex Hoffman, Deutschl. Fl. 2 (addenda), 11. 1796

Plants ca. 5--10 x 0.4--0.5(--1) mm, prostrate with apex frequently ascending, yellowish green to yellowish brown and deep green. Stems 0.11--0.15 mm in diameter, sparsely branching, orbicular in cross section, with homogenous inner cells, outer cells distinctly thickened, ventral surface brownish; rhizoids dense, in tangled fascicles, colorless to brownish, closely attached plants to substratum. Leaves subtransversely to obliquely inserted, appressed to the stem, imbricate, ends of lobes turned to  apex, orbicular to subquadrate, 0.5--0.7 x 0.7--1 mm, 2-lobed, subequal to unequal acute lobes divided by angular to crescentic sinus descending to 1/4\x leaf length; lobes entire, triangular, acute; cells of midleaf 5--6-angular with rounded angles, 25--35 x 22--27 \um, at base more large, to  ca. 40 x 35 \um; cuticle smooth, walls clearly thick, trigones concave and small; oil bodies (3--)6--12(--15) per cell, spheric ca. 5--8 \um in diameter to oval 5--8 x 6--10(--12) \um, granulate, colorless, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae common at apices of shoots, reddish yellow to reddish brown, commonly 4--5-angular in outline, with protruding angles, 24--38 x 24--28 \um, 2-celled. Sexual condition paroicous, very rarely with androecial subfloral innovations (autoicous). Androecia intercalary, male bracts 2--4(--5) pairs, transversely oval inflated in the base, 1--2-androus, stalk 1(--2)-seriate. Gynoecia terminal, female bracts in 1 pair, obovate to nearly orbicular, nearly entire, ca. 0.7--1.2 x 0.9 mm, divided to 1/3\x length into 2--3(--4) lobes, connate, bracteole present, lanceolate, sometimes with one additional teeth near base, connate with one of bracts to 2/3\x length. Perianth exerted to 2/3\x length, cylindric to cone-shaped, complicated at mouth, mouth lobulate and dentate with teeth to 1--2(--3) cells in length, generally 1-stratose. Capsule oval, exterior wall with nodular thickenings, interior wall with annular thickenings, reddish brown. Elaters ca. 8 \um in diameter. Spores 12--15 \um in diameter.

Fine-grained and sandy soil in various communities except Arctic deserts, mostly mesic conditions, sometimes growing among other liverworts in disturbed areas as well as on rocks covered by thin layer of mineral soil; 0--2700 m; Greenland; B.C., N.B., Nfld. and Labr., N.S., Ont., Que.; Alaska, Colo., Conn., Ga., Idaho, Ill, Ind., Kans., Ky., Maine, Mass., Mich., Minn., Miss., Mo., Nev., N.H., N.J., N.Y., N.C., Ohio, Okla., Oreg., Pa., R.I., S.C., Tenn., Vt., Va., Wash., W.Va., Wis.; Eurasia, Australia; Oceania (New Zealand).

20. Lophozia decolorans (Limpricht) Stephani, Spec. Hep. 2: 147. 1902

Jungermannia decolorans Limpricht, Jahresb. Schles. Gesell. Vaterl. Kult. 57: 116. 1880

Plants ca. 3--6 x 0.5--0.7 mm, prostrate to semi-immersed into substratum, yellow-brown with decolorate shoot apices. Stems 0.11--0.15 mm in diameter, sparsely branching, orbicular in cross section, with homogenous inner cells, outer cells distinctly thickened, ventral surface brownish; rhizoids dense, in tangled fascicles, colorless to brownish, closely attached plants to substratum. Leaves subtransversely inserted, appressed to the stem, imbricate, ends of lobes turned to  apex, widely ovoid-orbicular, 0.5--0.6 x 0.8--0.8 mm, 2-lobed, subequal to unequal acute lobes divided by U-shaped to crescentic sinus descending to 1/9--1/8\x leaf length; lobes entire, triangular, acute; cells of midleaf 5--6-angular with rounded angles, 23--30 x 12--23 \um; cuticle smooth, walls clearly thick at margins, but thinner in middle, trigones absent in margins to convex in midleaf; underleaves absent. Specialized asexual reproduction by gemmae common at apices of shoots, reddish, commonly 4--5-angular in outline, with slightly protruding angles, 24--26 x 19--25 \um, 1--2-celled. Sexual condition dioicous. Androecia intercalary, male bracts 3--5(--7) pairs, transversely oval inflated in the base, 1-androus, stalk 1(--2)-seriate. Gynoecia terminal, female bracts in 1 pair, obovate to nearly orbicular, nearly entire, ca. 0.7--0.8 x 1--1.3 mm, divided to 1/4\x length into 2--3 lobes, connate, bracteole present, lanceolate. Perianth exerted to 1/2\x length, cylindric to cone-shaped, complicated at mouth, mouth dentate with teeth to 1--2 cells in length, generally 1-stratose. Spores 12--15 \um in diameter.

Semi-immersed into soil over granite rocks (only one report for North America); 1600 m; B.C.; Eurasia; Tropical Africa

XXc. Lophozia subg. Sudeticae (Schljakov) Bakalin, Bryologist 114: 310. 2011

Lophozia sect. Sudeticae Schljakov, Pechen Mkhi Severa SSSR 3: 113. 1980

Plants ca. 5--25 x 0.7--2.2 mm, prostrate, brown to purplish brown, rarely deep brownish green. Stems 0.2--0.5 mm in diameter, sparsely branching, transverse section with or without microcellous layer to 1/2\x stem thickness, outer cells of stem in cross section thickened. Leaves oblique to subhorizontally inserted, more or less distant to subimbricate, spreading, 2(--3)-lobed; cuticle smooth to papillose; underleaves lanceolate or 2-lobed into 2 narrow lamina, filiform, frequently irregular and nearly absent. Specialized asexual reproduction by angular gemmae, with moderately thickened angles, colorless to brown. Sexual condition dioicous. Capsule oval, wall 3(--4)-stratose, Spores spheric, ca. 10 \um.

Species 2 (2 in the flora): North America, Eurasia.

1. Leaves 2--3(--4)-lobed, sinus descending to 1/3\x leaf length, underleaves commonly present . . . 22.Lophozia debiliformis

1. Leaves 2-lobed, sinus descending to 1/5\x leaf length, underleaves commonly absent . . . 21.Lophozia sudetica

21. Lophozia sudetica (Nees ex Huebener) Grolle, Trans. Brit. Bryol. Soc. 6: 262. 1971

Jungermannia sudetica Nees ex Huebener, Hepaticol. Germ., 142. 1834.

Plants ca. 5--20 x 1--2(--2.2) mm, prostrate to ascending or erect in dense patches, brownish to rusty brown, rarely with purple tinge or dark green (in shady places). Stems 0.2--0.4 mm in diameter, sometimes transversely oval in cross section, ca. 0.3 x 0.4 mm, sparsely branching, transverse section with microcellous layer to 1/3\x stem thickness or absent in southern phases, ventral surface brown to purple-black; rhizoids sparse to nearly absent, in indistinct fascicles, colorless to yellowish and brownish. Leaves commonly very obliquely inserted but rarely subtransverse, insertion line nearly straight, distant, spreading to appressed, concave to cup-shaped, turned to  apex or to dorsal side, orbicular to trapezoidal-rounded, (0.4--)0.6--1.2 x (0.5--)0.7--1.3 mm, 2-lobed, subequal lobes divided by crescentic to (rarely) angular sinus descending to (1/12--)1/8--1/5(--1/4)\x leaf length; lobes entire, acute, triangular; cells of midleaf 4--polygonal, 17--25(--27) x 13--22(--28) \um, in base to  40 \um; cuticle smooth to faintly verruculose, walls thin, trigones concave, triangular to convex; oil bodies (1--)2--6(--15) per cell, irregularly oval, 4--7 x 5--9 \um, rarely spheric, 4--7 \um in diameter, very finely papillose, grayish, non-biconcentric; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of distal leaf lobes, rusty-brown, commonly 4--5-angular in outline, 17--22 x 12--15 \um, (1--)2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 2(--3) pairs, similar to leaves but inflated in the base, 2(--3)-androus, antheridial stalk 1-seriate at base, but sometimes 2-seriate near spherule. Gynoecia terminal, innovations absent, female bracts in 1 pair, oval to ovoid, shallowly 2-lobed, entire, ca. 1.1 x 1 mm, divided to 1/7--1/5\x length into 2 lobes, connate, bracteole present, lanceolate, ca. 0.5--0.2 mm connate with one of bracts to 1/3\x length. Perianth exerted to 3/4\x length, cylindric to clavate and cylindric-fusiform, complicated at mouth, mouth denticulate to dentate with 2-celled teeth, last cell about 35 \um in length, generally 1-stratose. Seta ca. 3--5 mm, capsule oval, walls 3--4-stratose, exterior walls with nodular thickenings, interior walls with annular thickenings, reddish brown. Elaters 2-spiral, linear, reddish brown, ca. 100--125 x 7 \um. Spores ca. 10 \um in diameter.

Rocks outcrops, shaded soil, among mosses and other liverworts, common in both xeric and mesic conditions, more rarely hygrophilous, but frequent in snow-bed areas, tundra communities, sparse but frequent in some areas in boreal and temperate zones in rock crevices and soil on steep slope; 0--2500 m; Greenland; Alta., B.C., Nfld. and Labr., N.W.T., Nunavut, Ont., Que., Yukon; Alaska, Calif., Colo., Conn., Idaho, Maine, Mass., Mich., Minn., Mont., Nev., N.H., N.Y., Oreg., Utah, Vt., Wash., Wis., Wyo.; Eurasia.

22. Lophozia debiliformis R. M. Schuster & Damsholt, Phytologia 63(5): 326. 1987

Plants ca. 5--25 x 0.7--1(--2) mm, prostrate, nearly soft-textured, green-brownish to brown, apparently without purple tinge. Stems 0.25--0.4 mm in diameter, sometimes transversely oval in cross section, ca. 0.3 x 0.4 mm, sparsely branching, transverse section with homogenous cells without microcellous layer, ventral surface brownish; rhizoids sparse to nearly absent, in indistinct fascicles, colorless to yellowish and brownish. Leaves commonly obliquely inserted to rarely subtransverse, insertion line nearly straight or incurved  from oblique on ventral side to subtransverse on the dorsal, distant, spreading, concave to flattened and undulate, widely ovoid, orbicular to trapezoidal-rounded, (0.4--)0.6--1.2 x (0.5--)0.7--1.3 mm, 2--3(--4)-lobed, subequal or unequal lobes divided by angular or gibbous to U-shaped sinus descending to 1/5--1/3\x leaf length; lobes entire, acute, triangular; cells of midleaf 4--polygonal, 16--25(--27) x 16--22(--25) \um, in base to  30 \um; cuticle smooth, walls thin, trigones concave, triangular to convex; oil bodies (2--)3--5(--12) per cell, irregularly oval, 6--10 x 4--8 \um, rarely spheric, 4--6 \um in diameter, papillose to rarely smoothed botryoidal, grayish, non-biconcentric; underleaves present, lanceolate or 2-lobed into 2 narrow lamina, frequently irregular and nearly absent. Specialized asexual reproduction by gemmae in masses at apices of distal leaf lobes, rusty-brown to brownish white, commonly 4--6-angular in outline, 23--26 x 16--20 \um, (1--)2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 2--3(--4) pairs, similar to leaves but inflated in the base, 1(--2)-androus, antheridial stalk 1-seriate. Gynoecia terminal, innovations present, female bracts in 1 pair, oval to ovoid, deeply (2--)3-lobed, entire, but frequently with additional tooth near dorsal base, ca. 0.6 x 0.6 mm, divided to 1/4--2/5\x length into (2--)3 lobes, bracteole present, lanceolate, ca. 0.5--0.2 mm connate with one of bracts to 2/3\x length. Perianth exerted to 1/2\x length, obovate-clavate, complicated at mouth, mouth denticulate to dentate with 1--2-celled teeth, generally 1-stratose. Sporophyte unknown.

Snow-bed habitats, coastal cliffs, thin soil on shaded rocks, tundra zone; 0--1200 m; Greenland; Alaska; Eurasia.

XXd. LOPHOZIA subg. PROTOLOPHOZIA R. M. Schuster, Nova Hedwigia 15: 472. 1968

Plants ca. 5--25 x 0.7--2.2 mm, prostrate, brown to purplish-brown, rarely deep green-brownish. Stems 0.2--0.5 mm in diameter, sparsely branching, transverse section without microcellous layer, outer cells of stem in cross section thin. Leaves oblique to subhorizontally inserted, more or less distant to subimbricate, spreading, 2--3-lobed; cuticle smooth to papillose; underleaves lanceolate or 2-lobed into 2 narrow lamina, filiform, sometimes irregular. Specialized asexual reproduction not present in our species. Sexual condition paroicous. Capsule oval, walls 3(--4)-stratose, Spores spheric, 10--14 \um.

Species 11 (1 in the flora): North America; Central America; South America; Eurasia; Oceania; Australia, Africa.

23. Lophozia elongata Stephani, Spec. Hep. 2: 128. 1901

Plants ca. 5--20 x 1.1--1.5 mm, prostrate, slightly soft-textured, green to yellowish green and rarely green-brownish, apparently without purple tinge. Stems 0.2--0.3 mm in diameter, sometimes transversely oval in cross section, ca. 0.2 x 0.3 mm, sparsely branching, transverse section with homogenous cells without microcellous layer, ventral surface indistinguishable from the dorsal; rhizoids sparse to nearly absent, in indistinct fascicles, nearly colorless. Leaves commonly obliquely inserted to rarely subtransverse, insertion line nearly straight or incurved from oblique in ventral side to subtransverse in the dorsal, distant, spreading, concave to flattened and undulate, widely ovoid, orbicular, trapezoidal-rounded or obtrapezoidal, (0.4--)0.6--0.7 x (0.5--)0.6--0.7 mm, 2--3(--4)-lobed, subequal or unequal lobes divided by angular or gibbous to U-shaped sinus descending to 1/5--1/3\x leaf length, additional teeth frequently present near leaf base; lobes entire, acute, triangular; cells of midleaf 4-polygonal, 23--28 x 21--25 \um; cuticle smooth, walls thin to slightly thickened, trigones concave to nearly absent; oil bodies numerous, 10--42 per cell, small, spheric, 2--4.5 \um in diameter, finely papillose; underleaves present, lanceolate, 2-lobed into 2 narrow lamina or reduced to slime papilla. Specialized asexual reproduction absent. Sexual condition paroicous. Androecia intercalary, male bracts 2--3(--4) pairs, similar to leaves but inflated in the base, additional tooth near antical base commonly present, 1-androus, antheridial stalk 1-seriate. Gynoecia terminal, female bracts in 1 pair, oval to ovoid, deeply (2--)3-lobed, nearly entire, but sometimes with additional teeth, ca. 0.6 x 0.6 mm, divided to 1/4--2/5\x length into (2--)3 lobes, bracteole present, deeply 2-lobate, ca. 0.5--0.2 mm connate with bracts to 1/2\x length. Perianth exerted to 3/4\x length, obovate-clavate to cylindric, complicated at mouth, mouth denticulate to dentate with 1--2-celled teeth, generally 1-stratose. Elaters ca. 6 \um thick. Spores 10--14 \um in diameter.

Permanently damp peaty slopes, wet moss mats in moist tundra, among or over mosses and other liverworts in dense patches; elevation unknown; Greenland; Alta.; n Eurasia.

XXe. LOPHOZIA subg. OBTUSIFOLIUM (H. Buch) Bakalin

Barbilophozia subg. Obtusifolium H. Buch, Mem Soc. F. Fl. Fenn. 17: 289. 1942

Plants ca. 20--50 x 2--3.5 mm, prostrate, greenish brown, brownish to brownish yellow, rarely with purple tinge or dark green in shady places. Stems 0.3--0.5 mm in diameter, sparsely branching, transverse section commonly without microcellous layer, but sometimes with exception of 2--3 layers of cells smaller than inner ones, outer cells in stem cross section thin; ventral side of the stem nearly the same color with the dorsal. Leaves subhorizontally inserted, more or less distant, spreading, flattened, 2(--3)-lobed, cells with many (to 50 per cell) small, spheric oil-bodies; cuticle distinctly papillose; underleaves commonly absent, rarely present, ciliate or lacinulate, 4 cells in length and 1--2(--3) cells wide, easily deciduous. Specialized asexual reproduction by angular gemmae, those rarely present, green, with moderate thickened angles. Sexual condition dioicous. Capsule oval, walls (4--)5-stratose, Spores spheric, 11--14 \um.

Species 1 (1 in the flora): North America, Europe, Asia, Africa.

24. Lophozia obtusa (Lindberg) A. Evans, Proc. Washington Acad. Sci. 2: 303. 1900

Jungermannia obtusa Lindberg, Musci Scand., 7. 1879

Plants ca. 20--50 x 2--3.5 mm, prostrate or epiphyte on sedges or other boggy grasses, greenish brown, brownish to brownish yellow, rarely with purple tinge or dark green (in shady places). Stems 0.3--0.45 mm diameter, sometimes transversely oval in transverse section (ca. 0.25 mm in height), sparsely branching, transverse section commonly without microcellous layer, but occasionally with 2--3 layers of cells smaller than inner ones located above ventral cortex, ventral surface brown to reddish brown; rhizoids common sometimes dense, in indistinct fascicles, colorless to yellowish and brownish. Leaves commonly very obliquely inserted to nearly horizontally, insertion line nearly straight, distant, spreading, flattened, lingulate to orbicular, 1--1.2 x 1.2--1.4 mm, 2-lobed, subequal lobes divided by gibbous sinus descending to 1/5(--1/4)\x leaf length; lobes entire, rounded to obtuse; cells of midleaf 4--polygonal, 32--38 x 21--25 \um, in base to  50 x 38 \um; cuticle almost always distinctly striolate papillose through, especially obvious at the base, walls thin, trigones concave to triangular; oil bodies minute, 15--50 per cell, spheric, 2--3 \um in diameter, nearly smooth; underleaves commonly absent, rarely present, ciliate or lacinulate, 4 cells in length and 1--2(--3) cells wide, easily deciduous. Specialized asexual reproduction by gemmae at shoot apices, rarely present, pale green to brownish, commonly with 4--5-angular in outline, 17--22 x 17--20 \um, 2-celled, with slightly thickened angles. Sexual condition dioicous. Androecia intercalary, male bracts 6--12 pairs, spicate, 2--3-androus, antheridial stalk 1-seriate. Gynoecia terminal, innovations absent, female bracts in 1 pair, oval to ovoid, 2--4-lobed, entire, ca. 1.1 x 1 mm, divided to 1/7--1/5\x length, bracteole present, 2-lobed. Perianth exerted to 3/4\x length, cylindric, ellipsiodal to clavate, complicated at mouth, mouth denticulate to dentate. Seta ca. 3--5 mm, capsule oval, wall 5-stratose, exterior wall with nodular thickenings, interior walls with annular thickenings, reddish brown. Elaters ca. 7--9 \um wide. Spores ca. 11--14 \um in diameter.

Isolated stems (never in pure mats) among mosses, wet meadows, green-moss bogs, sometimes on moist slopes or waterways; 0--2150 m; Greenland; Alta., B.C., Nfld. and Labr. (Nfld.), N.W.T., Nunavut, Que., Yukon; Alaska, Ariz., Calif., Colo., Idaho, Maine, Mich., Mont., Oreg., Wash., Wyo.