BFNA Title: Tritomaria
Author: W. S. Hong
Date: March 21, 2002
Edit Level: R Brum+
Version: 1

Bryophyte Flora of North America, Provisional Publication
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Tritomaria, Jungermanniaceae

TRITOMARIA Loeske, Hedwigia 49: 13. 1909

Won Shic Hong

Plants ca. 0.5--5 cm ´ 1--4 mm, prostrate to ascending. Stem transverse section with 1--3 cortical cells and 15--25 medullary cells; rhizoids numerous in dense mats, colorless to pale brown. Leaves subtransversely-obliquely inserted, 2--4-lobed, asymmetrical (except T. polita); lobes triangular, unequal, margins entire; cell walls collenchymatous, trigones bulging; oil bodies 3--8 per cell, finely granular, spherical to ovoid; underleaves absent. Sexual condition dioicous. Specialized asexual reproduction by gemmae. Androecia intercalary, antheridial stalk 1--2-seriate, female bracts leaflike, 3--4-lobed, margins entire-dentate, bracteoles usually absent. Perianth cylindrical-ovoid, plicate, mouth entire or ciliate. Capsule oval, walls 3--5-stratose, exterior wall with nodular and interior wall with semiannular bands. Elaters ca. 6--10 m m diam. Spores ca. 10--16 m m.

Species 8 (6 in the flora): North America, Europe, Asia, Africa.

SELECTED REFERENCES

Schuster, R. M. 1969. Tritomaria Schiffn. In: R. M. Schuster. 1966--1993. The Hepaticae and Anthocerotae of North America. New York. Vol. 2, pp. 638--704. Schuster, R. M. 1984. Evolution, Phylogeny and Classification of the Hepaticae. In: R. M. Schuster, ed. 1984. New Manual of Bryology. Vol. 2, pp. 892--1070. Nichinan.

1. Leaves more or less symmetrical, equally 2--4-lobed with blunt to rounded apices; perianth mouth entire or subentire . . . . 4. Tritomaria polita

1. Leaves asymmetrical, unequally 2--3-lobed with acute or acuminate apices; perianth mouth dentate.

2. Leaves complicate to explanate, wider than long; gemmae rare-absent, yellowish-reddish brown.

3. Plants 1--1.5 mm wide, pellucid; leaves approaching symmetrical form, loosely complex, trigones coarsely nodose; gemmae angulate; perianth mouth teeth 1--2 cells long . . . . . 3. Tritomaria heterophylla

3. Plants 2--3.5 mm wide, opaque; leaves asymmetrical, explanate, trigones moderate-bulging, but never nodose; gemmae generally absent; perianth mouth teeth 4--5 cells long . . . . 5. Tritomaria quinquedentata

2. Leaves canaliculate-complanate, longer than wide; gemmae abundant, reddish brown to rust red.

4. Leaves usually 3-lobed, lobes acute to subacute; trigones large; cilia of perianth mouth 1--4 cells long; gemmae reddish-brown . . . . . 6. Tritomaria scitula

4. Leaves asymmetrically 2--3-lobed, lobes acute or acuminate; trigones small; cilia of perianth mouth 5--6 cells long; gemmae rust-red.

5. Gemmae smooth, ovoid-elliptical; leaf cells 8--14 ´ 10--20 m m in midleaf; cell walls thick, trigones not bulging . . . . 1. Tritomaria exsecta

5. Gemmae angulate, polygonal; leaf cells 18--22 ´ 20--35 m m in midleaf; cell walls thin, trigones bulging . . . . 2. Tritomaria exsectiformis

1. Tritomaria exsecta (Schrader) Loeske Hedwigia 49: 13. 1909

Jungermannia exsecta Schrader, Syst. Samml. Cryptogamisher Gewasche 2: 5. 1797

Plants ca. 0.5-- 2 cm ´ 1--2 mm, ascending, pale green to brownish green. Stems 0.2--0.3 mm diam., sparsely branching, transverse section with 2--3 cortical cells and 12--18 medullary cells; rhizoids numerous, colorless-pale brownish. Leaves succubous, distant to imbricate, conduplicate-concave, ovate to oblong-ovate, 0.8--1.2 ´ 0.5--0.8 mm, unequally 2--3-lobed (median lobes often absent); antical lobes subulate, more weakly arched than postical lobe, sinus between middle lobes deeper (to 0.3 of the leaf length), margins entire, apex acuminate to acute; postical lobes ovate, more strongly arched than antical lobes, sinus between middle lobes very shallow (less than 0.1 of the leaf length), margins entire, apex acuminate to acute; cells rounded-rectangular, marginal cells 8--12 ´ 10--15 m m, median cells 8--14 ´ 10--20 m m , basal cells 10--15 ´ 15--30 m m, cuticle verruculose, walls thick, trigones small; oil bodies filling the lumen, 2--8 per cell, spherical to ovoid, 2--4 ´ 4--5 m m, finely granular, grayish; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of shoots, rust-red, elliptical, 15--22 ´ 8--12 m m, 2-celled, smooth. Sexual condition dioicous. Androecia terminal, often with gemmae at tips, male bracts 4--10 pairs, similar to leaves, with antheridia, 1--2 androus, antheridial stalk 1-seriate. Gynoecia terminal, female bracts in few pairs, rounded -quadrate to broadly ovate, 3--5 lobed, equal or slightly smaller than leaves. Perianth cylindrical-obovate, 4--6-plicate, mouth ciliated with teeth 2--6 cells long. Seta 5 mm, capsule ovoid, walls 3-stratose, exterior walls with nodular thickenings, interior walls with annular thickenings, reddish brown. Elaters ca. 8 m m in diam., 2-spiral, reddish brown. Spores 9--12 m m, papillose, dark brownish.

Humic soil, humus on rocks, trunks and branches, especially Alnus and Malus in the West; 0--2000 m; Alta., B.C., N.B., Nfld., N.W.T., N.S., Ont., Que.; Alaska, Colo., Conn., Ga., Idaho, Iowa, Ky., Maine, Mass., Mich., Minn., Mont., N.H., N.Y., N.C., Ohio, S.C., Vt., Va., Wash., W.Va.; Mexico; e Africa.

In the East T. exsecta is frequently associated with Anastrophyllum minutum, A. michauxii, Bazzania trilobata, Diplophyllum apiculatum, Frullania asagrayana, Harpalejeunea ovata, Herbertus aduncus, Jamesoniella autumnalis, Lejeunea cavifolia, L. ulicina, Metzgeria crassipilis, Radula obconica, R. tenax, and Scapania nemorosa. In the West it is associated with Blepharostoma trichophyllum, Douinia ovata, Frullania nisquallensis, Scapania bolanderi, and S. umbrosa.

2. Tritomaria exsectiformis (Breidler) Loeske

Jungermannia exsectiformis Breidler, Mitt. Naturwiss. Vereines Steiermark 30: 321. 1894

Plants ca. 0.5--3 cm ´ 1.5--2 mm, ascending, green to brownish. Stems 0.2--0.3 mm diam., sparsely branching, transverse section with 2--3 cortical cells and 12--18 medullary cells; rhizoids colorless to brownish. Leaves succubous, contiguous to imbricate, conduplicate-concave, ovate to oblong-ovate, 0.8--1.2 ´ 0.5--0.8 mm, unequally 2--3-lobed (median lobes often absent); antical lobes smaller than postical lobes, apex acuminate to acute, margins entire; cells rounded-rectangular, marginal cells 14--18 ´ 18--22 m m, median cells 14--22 ´ 20--35 m m, basal cells 20--25 ´ 28--40 m m, cuticle strongly verruculose, walls thin, trigones strongly bulging; oil bodies 5--12 per cell, spherical to ovoid, 3--5 ´ 4--6 m m, granular, grayish; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of shoots, rust-red or bright red, polygonal to pyriform, 12--25 ´ 10--25 m m, 2-celled. Sexual condition dioicous. Androecia terminal, often with gemmae at tips, male bracts similar to leaves, 3-lobed, with antheridia, 1--2 androus, antheridial stalk 1-seriate. Gynoecia terminal, female bracts larger than leaves, ovate, 3--4-lobed. Perianth free from the bracts, cylindrical, 4--6 plicate, mouth constricted, lobulate with teeth 2--5 cells long. Capsule ovoid, walls 2--4-stratose, exterior walls with nodular thickenings, interior walls with semiannular thickenings, reddish-brown. Elaters ca. 7--10 m m diam., 2-spiral, reddish-brown. Spores 9--12 m m, papillose.

Subspecies 2 (2 in the flora): widely distributed across the northern part of the floral range.

1. Leaf cells 18--22 ´ 20--35 m m in midleaf; gemmae irregularly polygonal to pyriform, rust-red, 15--25 ´ 12--20 m m; widespread . . . . 2a. Tritomaria exsectiformis subsp. exsectiformis

1. Leaf cells 14--16 ´ 20--25 m m in midleaf; gemmae angulate, bright reddish, 12--25 ´ 10--25 m m; arctic . . . . 2b. Tritomaria exsectiformis subsp. arctica

2a. Tritomaria exsectiformis (Breidler) Loeske subsp. exsectiformis

Leaf cells 18--22 ´ 20--35 m m in midleaf; gemmae irregularly polygonal to pyriform, rust-red, 15--25 ´ 12--20 m m.

Creek banks, decayed wood, and humus over decayed wood in forests; 0--2300 m; Greenland; Alta., B.C., Nfld., N.W.T., N.S., Nun., Ont., Que., Yukon; Alaska, Colo., Conn., Idaho, Iowa, Maine, Mass., Mich., Minn., Mont., N.H., N.Y., N.C., Pa., Vt., Wash., Wis., Wyo.; Europe; Asia.

In the East it is frequently associated with Bazzania trilobata, Lophozia ascendens, L. longidens, and Tritomaria exsecta. In the West it is associated with Blepharostoma trichophyllum, Cephalozia lunulifolia, Gymnomitrion coralloides, Lepidozia reptans, Lophozia incisa, L. ventricosa, and Ptilidium pulcherrimum.

2b. Tritomaria exsectiformis subsp. arctica R. M. Schuster, Hep. Anthoc. N. Amer. 2: 661. 1969

Leaf cells 14--16 ´ 20--25 m m in midleaf; gemmae angulate, bright reddish, 12--25 ´ 10--25 m m. Androecia, gynoecia and sporophyte unknown.

Peaty soil with acidic rocks; 50--80 m; endemic to w Greenland.

This North American endemic is associated with Anastrophyllum minutum, A. sphenoloboides, Cephalozia bicuspidata, Cephaloziella divaricata, Lophozia opacifolia, Scapania lingulata, and Tritomaria quinquedentata.

3. Tritomaria heterophylla R.M. Schuster, Canad. Jour. Bot. 36: 272. 1958

Plants ca. 0.8--1.5 cm ´ 0.7--1.5 mm, ascending, brownish-purplish to brown. Stems 0.3--0.4 mm in diam., sparsely branching, transverse section with cortical and medullary cells; rhizoids numerous, colorless-pale brownish. Leaves succubous, imbricate, loosely complicate, transversely oblong, 0.7--0.9 ´ 0.8--1.2 mm, subequally 3-lobed, lobes ovate-triangular, sinus 0.2-0.3 of leaf length, apex acuminate to acute, margins entire or with scattered sharp teeth; cells rounded rectangular, marginal cells 20--25 m m, median cells 20--25 ´ 25--30 m m, basal cells 25--30 ´ 30--40 m m, cuticle weakly verruculose, strongly collenchymatous, trigones bulging; oil bodies 2--10 per cell, spherical to elliptical, 6--7 ´ 9--13 m m, finely papillose, grayish; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of juvenile leaves, reddish-brown, angular, 25--35 ´ 18--30 m m, 1--2 celled, smooth. Sexual condition dioicous. Male bracts similar to leaves, antheridia orange-yellow, antheridial stalk 1-seriate. Gynoecia terminal, female bracts larger than leaves, 2--5-lobed, lobes broadly ovate-triangular with entire margins. Perianth ovoid, plicate, mouth ciliated with teeth 2 cells long. Sporophyte unknown.

Peaty soil between rock clefts; 0--250 m; e, w, and n Greenland; Nun.

This species is superficially similar to Tritomaria scitula and T. quinquedentata, and is frequently associated with Aneura pinguis, Antheria juratzkana, Arnellia fennica, Blepharostoma trichophyllum subsp. brevirete, Cephalozia bicuspidata, C. pleniceps, Cephaloziella arctica, Cryptocolea imbricata, Gymnomitrion concinnatum, Jungermannia sphaerocarpa, Odontoschisma macounii, Plagiochila arctica, Prasanthus suecicus, Scapania gymnostomophila, and Tritomaria quinquedentata.

4. Tritomaria polita (Nees) Jørgensen, Bergens Mus. Aarbok 7: 4. 1921

Jungermannia polita Nees, Naturg. Eur. Leberm. 2: 9. 1836

Plants ca. 1--1.5 cm ´ 2--3 mm, ascending, yellowish-green to reddish-brown. Stems 0.2--0.5 mm in diam., sparsely branching with Frullania-type, Radula-type and lateral-intercalary; transverse section with 1--3 elongated cortical cells and 10--16 medullary cells; rhizoids moderately numerous, colorless. Leaves transversely inserted, symmetrical, variously wider or narrower than long, broadly quadrate, ca. 1.7--2 ´ 1.4--1.7 mm, 3 or 2--4-lobed, sinus very shallow (0.1--0.25 of the leaf length), apex obtuse to rounded, margins entire; cells rounded-polygonal, marginal cells 15--25 ´ 25--35 m m, median cells 25--30 ´ 45--50 m m, basal cells 35--45 ´ 70--100 m m, cuticle weakly verruculose, walls thin, trigones large; oil bodies 2--12 per cell, spherical to ellipsoidal, ca. 5--10 m m, granular, grayish; underleaves absent. Specialized asexual reproduction by gemmae in masses at tips of leaf-lobes, yellow-brown to purplish, smooth or angulate, elliptical, ca. 28 m m, 1--2-celled. Sexual condition dioicous. Androecia terminal, male bracts, several pairs, similar to leaves, 3-lobed, lobes undulate. Seta 1--2 cm, capsule ovoid, exterior walls with nodular thickenings, interior walls with semiannular thickenings. Elaters ca. 8 m m in diam., 2-spiral, reddish-brown. Spores ca. 15 m m, reddish-brown.

Subspecies 2 (2 in the flora): North America, Europe, Asia.

1. Leaves uniformly 3-lobed with angulate sinuses, longer than wide; oil bodies 2--7 per cell; gemmae brown to purplish, angulate; female bracts 3-lobed; widespread . . . . . 4a. Tritomaria polita subsp. polita

1. Leaves 2--4-lobed with curved sinuses, wider than long; oil bodies 8--12 per cell; gemmae usually absent, if present yellow brown, smooth; female bracts 2--4-lobed; sub-Arctic and Arctic . . . . 4b. Tritomaria polita subsp. polymorpha

4a. Tritomaria polita (Nees) Jørgensen subsp. polita

Leaves uniformly 3-lobed with angulate sinuses, longer than wide; oil bodies 2--7 per cell; gemmae brown to purplish, angulate; female bracts 3-lobed.

Exposed boulders, stream banks, soil on cliffs in forests, and seepage slopes in the tundra.; 500--1100 m; Greenland; Alta., B.C., Nfld. (Labrador), Nun., Que.; Alaska, Colo., Mont., Wash.; Europe; Asia.

It is frequently associated with Barbilophozia kunzeana, Blepharostoma trichophyllum, Calypogeia azurea, Cephalozia bicuspidata, Diplophyllum albicans, Harpanthus flotovianus, Jungermannia atrovirens, Leiocolea heterocolpos, Lophozia guttulata, L. incisa, Pleuroclada albescens, and Tritomaria exsectiformis.

4b. Tritomaria polita subsp. polymorpha R. M. Schuster, Hep. Anthoc. N. Amer. 2: 700. 1969

Leaves 2--4-lobed with curved sinuses, wider than long; oil bodies 8--12 per cell; gemmae usually absent, if present yellow brown, smooth; female bracts 2--4-lobed. Sporophyte unknown.

Banks of small streams; 0--50 m; Greenland.

This North American endemic taxon is reported only from Greenland (R. M. Schuster 1988). It is frequently associated with Blepharostoma trichophyllum subsp. brevirete, Cephalozia pleniceps, Leiocolea heterocolpos, Lophozia gillmanii, Odontoschisma elongatum, Scapania brevicaulis, and Tritomaria quinquedentata. R. Grolle and D. G. Long (2000) treated this taxon as T. polita.

5. Tritomaria quinquedentata (Hudson) H. Buch, Mem. Soc. Fauna Fl. Fenn. 8: 270. 1932

Jungermannia quinquedentata Hudson, Fl. Angl. ed. 1: 433. 1762; Lophozia quinquedentata (Hudson) Cogniaux

Plants ca. 1.5--6 cm ´ 2-4 mm, ascending , green to yellowish-brown. Stems 0.3-0.45 mm in diam., sparsely branching, transverse section with 2--3 cortical cells and 10--14 medullary cells; rhizoids numerous, colorless. Leaves succubous, imbricate, ovate-reniform, 0.8--1.4 ´ 1--1.6 mm, unequally 3-lobed, shallow sinus (0.1--0.2 of leaf length), lobes broadly ovate, apex acute to apiculate, margins entire; cells rounded polygonal, marginal cells ca. 12--18 ´ 20--23 m m, median cells ca. 18--24 ´ 21--28 m m, basal cells ca. 20--28 ´ 28--38 m m, cuticle verruculose, walls thin, trigones distinct; oil bodies 2--10 per cell, spherical to ovoid, 4--7 ´ 5--10 m m, finely granular, grayish; underleaves absent. Specialized asexual reproduction by gemmae, rare, at tips of leaf lobes, yellowish-brown, polygonal, ca. 15--20 m m, 1--2-celled, smooth. Sexual condition dioicous. Androecia terminal, male bracts 4--12 pairs, similar to leaves, 2--3-androus, antheridial stalk 2-seriate. Gynoecia terminal, female bracts larger than leaves, 3--5-lobed, lobes acute. Perianths cylindrical-obovoid, plicate, mouth ciliated with teeth 4--5 cells long. Seta 2 cm, capsule oblong-ovoid, walls 4--5-stratose, yellowish or reddish-brown, exterior walls with nodular thickenings, interior walls with semiannular thickenings. Elaters ca. 6--7 m m in diam., 2-spiral, reddish-brown. Spores 12--15 m m, verruculose, yellowish-brown.

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

1. Cells smaller, 18--24 ´ 21--28 m m in midleaf, trigones distinct; oil bodies 2--10 per cell; widespread . . . . 5a. Tritomaria quinquedentata var. quinquedentata

1. Cells larger, 24--32 ´ 30--42 m m in midleaf, trigones bulging; oil bodies 7--20 per cell; sub-Arctic and Arctic.

2. Cells 25--30 ´ 30--42 m m midleaf; oil bodies 10--20 per cell . . . . .5b. Tritomaria quinquedentata var. grandiretis

2. Cells 24--26 ´ 32--45 m m in midleaf; oil bodies 5--15 per cell . . . . 5c. Tritomaria quinquedentata var. turgida

5a. Tritomaria quinquedentata (Hudson) H. Buch var. quinquedentata

Plants green to yellowish-brown or pale brown. Leaf cells 18--24 ´ 21--28 m m in midleaf, trigones distinct but never bulging, oil bodies 2--10 per cell.

Wet humus over boulders, shaded cliffs, soil over exposed rock surface, decaying branches at the fringes of spray zones, and among heather on slopes; 0--1240 m; e, nw and sw Greenland; Alta., B.C., Man., N.B., Nfld. (Labrador), N.W.T., N.S., Nun., Ont., Que., Sask., Yukon; Alaska, Colo., Conn., Maine, Mich., Minn., Mont., N.H., N.Y., Oreg., Vt., Wash., Wis.; Europe; Asia.

This subspecies is frequently associated with Anastrophyllum minutum, Barbilophozia attenuata, B. barbata, Tetralophozia setiformis, Diplophyllum taxifolium, Geocalyx graveolens, Gymnomitrion coralloides, Leiocolpos heterocolpos, Lophozia sudetica, Mylia taylori, Odontoschisma macounii, Plagiochila asplenioides, Ptilidium ciliare, Scapania americana, S. mucronata, and S. nemorosa.

5b. Tritomaria quinquedentata var. grandiretis H. Buch & S. W. Arnell, Svensk Bot. Tidskr. 44 (1): 84. 1950

Plants pale and subpellucid. Leaf cells 25--32 ´ 30--42 m m in midleaf, trigones distinct, oil bodies 10--20 per median cell.

Acidic sites over peaty grounds near small stream; 0--500 m; nw and w Greenland.

This variety is restricted to Greenland (R. M. Schuster 1958, 1988; R. M. Schuster and K. Dansholt 1974), where it is frequently associated with Anastrophyllum minutum, Barbilophozia binsteadii, B. hatcheri, B. kunzeana, B. quadriloba, Lophozia ventricosa, Tritomaria quinquedentata subsp. quinquedentata, and Ptilidium ciliare.

5c. Jungermannia quinquedentata var. turgida Lindberg in Lindberg & Arnell, Musci Asiae Borealis I. Kgl. Sv. Vetensk. Akad. Handl. 23 (5): 59. 1889

Tritomaria quinquedentata subsp. turgida (Lindberg) Damsholt, Bryologist 85 (1): 96. 1982

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Plants uniformly strongly brownish. Leaf cells 24--26 ´ 30--45 m m in midleaf, trigones coarse and bulging; oil bodies 5--15 per cell.

Arctic and sub-Arctic fens, bogs, and on hillsides. It is frequently associated with Antheria juratzkana, Barbilophozia binsteadii, B. kunzeana, Blepharostoma trichophyllum subsp. brevirete, Cephaloziella arctica, Lophozia ventricosa, and Odontoschisma macounii; 0--3400 m; e and nw Greenland; Yukon; Alaska; Asia (Siberia).

The plants grow to 4 mm in width and the perianth is strongly ciliate-dentate with teeth 4--6 cells long, and 2--4-stratose at base in contrast to 1--3-stratose in subsp. quinquedentata.

6. Tritomaria scitula (T. Taylor) E. H. Jörgensen, Bergens Mus.Aarbok, Naturv. 7: 9. 1921

Jungermannia scitula T. Taylor London Jour. Bot. 5: 274. 1846

Plants ca. 0-5--1.5 cm ´ 1.5--2 mm, prostrate-ascending, green-brownish. Stems 0.33--0.35 mm in diam., sparsely branching, transverse section with 1--2 cortical cells and 12--16 medullary cells; rhizoids numerous, colorless. Leaves succubous, imbricate, canaliculate, ovate-oblong, 0.7--1 ´ 0.6--0.8 mm, unequally 3-lobed (occasionally 2--4-lobed); lobes acute to subacute, sinus between lobes shallow (0.2--0.3 of the leaf length), apex rounded to truncated, margins entire; cells subquadrate, marginal cells 16--22 ´ 24--30 m m, median cells 22--28 ´ 26--36 m m, basal cells 25--30 ´ 35--55 m m, cuticle verruculose, walls thin, trigones large (weakly bulging); oil bodies 4--10 per cell, spherical to ovoid, 5--8 ´ 5--10 m m, papillose; underleaves absent. Specialized asexual reproduction by gemmae in masses at apices of shoots and tips of leaf lobes, reddish-brown, polygonal, 16--24 ´ 20--30 ´ 15--25 m m, 1--2-celled, smooth. Sexual condition dioicous. Androecia terminal, male bracts similar to leaves, 2-androus, antheridial stalk 1-seriate. Gynoecia terminal, female bracts larger than leaves, broadly oblong, 3--4-lobed with acute sinuses. Perianths cylindrical, 4--5- plicate, mouth ciliated with teeth 2--5 cells long. Capsule ovoid, wall 4-stratose, both outer and interior walls with nodular thickenings, yellowish-brown. Elaters ca. 9 m m in diam., 2-spiral, brownish. Spores 14--16 m m, papillose, brownish.

With other bryophytes on calcareous soil and decayed wood; 0--3500 m; e, w, nw Greenland; Alta., B.C., N.B., N.W.T., Nun., Que., Yukon; Alaska, Mich., Minn., Mont., Wis.; Europe.

The sporophyte of this species is very rare. Tritomaria scitula is frequently associated with Anastrophyllum minutum, Barbilophozia binsteadii, B. floerkei, B. lycopodioides, B. quadriloba, Blepharostoma trichophyllum subsp. brevirete, Cephalozia pleniceps, Jungermannia sphaerocarpa, Leiocolea heterocolpos, Odontoschisma macounii, Plagiochila asplenioides, Tritomaria exsectiformis, T. quinquidentata, Scapania cuspiduligera, and S. mucronata.

OTHER REFERENCES

Grolle, R. and D. G. Long. 2000. An annotated check-list of the Hepa;ticae and Anthocerotae of Europe and Macaronesia. J. Bryol. 22: 103--140.

Schuster, R.M. 1958. Notes on Nearctic Hepaticae XIII The genus Tritomaria (Lophoziaceae) in Arctic Canada. Canad. Jour. Bot. 36: 269--288.

Schuster, R. M. 1988. The Hepaticae of South Greenland. Nova Hedwigia. Berlin. Vol. 92.

Schuster, R.M. & K. Damsholt. 1974. The Hepaticae of west Greenland from ca. 60° N to 72° N. Copenhagen.