BFNA Title:Gymnomitriaceae
Date: Jan. 24, 2020
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Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden
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Plants forming mats or turfs. Branches sometimes intercalary from sides of stem, sometimes replacing ventral half of a leaf; with or without flagella.  Leaves alternate, succubous, plane or concave, simple or shallowly 2-lobed with a narrow sinus, entire or finely serrulate. Underleaves small or absent, unlobed.  Rhizoids scattered over ventral stem surface. Specialized asexual reproduction absent.  Gynoecium terminal on an ordinary leafy branch, with or without subfloral branches. Perianth small or absent, included in bracts and completely hidden by them,  when present cylindrical, mouth narrow, often with a well-developed perigynium.


Genera 9 (4 in the flora), species ca. 70 (35 in the flora): worldwide including Antarctica.


Although excluded from the Gymnomitriaceae by J. Váňa et al. (2010). Nardia was added to the family by Váňa et al. (2014) and Söderström et al. (2016) based on molecular phylogenetic studies of various authors.


1, Leaves succubous, never transverse, undivided or bilobed, not interlocking dorsally (insertions not extending across the stem midline). Underleaves always present, mostly small, lanceolate. Gametangia on leading axes. Sporophytes enclosed by a shoot calyptra, perianth and perigynium (erect, of Isotachis-type or short pendent, of Nardia geoscyphus-type) always present. Capsules subsphaeroidal to shortly ellipsoidal, innermost wall cells with semiannular thickenings.  …3. Nardia, p. XXX



SELECTED REFERENCES  Söderström, L, A. Hagborg, M. von Konrat et al. 2016. World checklist of hornworts and liverworts. PhytoKeys 59: 1–828.   Váňa, J., L. Söderström, A. Hagborg, & M. von Konrat. 2014. Notes on early land plants today. 60. Circumscription of Gymnomitriaceae (Marchantiophyta). Phytotaxa 183(4): 287–289.  Váňa, J., L. Söderström, A. Hagborg, M. J. von Konrat and J. J. Engel. 2010. Early Land Plants Today: Taxonomy, systematics and nomenclature of Gymnomitriaceae. Phytotaxa 11: 1--80.


1. GYMNOMITRION Corda, in Opiz, Naturalientausch 12: 651. Sep. 1829, nom. cons. [Greek gymnos, naked, without, and mitrion, small headgear, crown; alluding to the absence of a perianth]


Yuriy S. Mamontov


Plants pale to yellow green or yellowish or chestnut-red or brownish or reddish black in color. Branching intercalary, usually from the older shoot sectors; the branches usually dimorphic, some ascending and leafy, others descending, reduced-leaved and rhizoidous, stolon-like or flagelliform; subgynoecial innovations usually frequent.  Rhizoids hyaline, yellowish to purplish, chiefly from stolons and older shoot sectors.  Stem soft to rigid, in cross section lacking a hyalodermis, with slightly to very thick-walled cortical and intracortical cells grading gradually into the collenchymatous, firm, pale medulla.  Leaves transversely to subtransversely inserted (slightly succubous); not decurrent or decurrent dorsally and ventrally; imbricate, erect-appressed or distant and patent to almost at right angle to the stem; not or slightly to distinctly increasing in size towards shoot apex; broadly ovate or oval or obovate, little constricted basally; concave (often strongly so); 2-lobed 0.1--0.45 of the length, or emarginate (rarely unlobed); the leaf tips and margins sometimes narrowly to extensively decolorate or not decolorate; plane or revolute, even in the sinuses; entire or sometimes erose with age, or with a single row of elongate, hyaline cells, projecting as crenulations or teeth.  Leaf cells collenchymatous, often with bulging trigones, rather small. Cuticle delicately papillose or smooth.  Oil bodies 1--5 per cell, usually ellipsoidal, rarely spherical or ovoid, granular, colorless, 2--8(--12) x 2--5 \um.  Underleaves absent or vestigial.  Sexual condition paroicous, autoicous or dioicous.  Androecia terminal or intercalary, composed of several pairs of imbricate bracts; each bract subtending (1--)2--3 antheridia.  Gynoecia terminal on main stems; bracts often in several imbricate pairs, much larger and often wider than vegetative leaves, sometimes with more or less lacerate, reduced innermost bracts; bracteole lacking.  Perianth lacking or vestigial or rather large, inflated, plicate.  Shoot calyptra often distinct, almost always replacing the perianth.  Sporophyte seta of numerous cell rows, often short.  Capsule spherical, usually reddish or dark brown. Capsule wall 2--3-stratose; all layers with nodular thickenings.  Spores mostly 8--16 \um, almost smooth to finely verruculose. Elaters 2--4-spiral.


Species 36 (11 in the flora): North America; Central America; South America; Arctic; Europe; Asia; Africa; Atlantic Islands; Indian Ocean Islands; Pacific Islands; Antarctic.


The taxonomy of Gymnomitrion follows B. Crandall-Stotler et al. (2009) and B. Crandall-Stotler and R. E. Stotler (2017). According to B. Shaw et al. (2015) the genus Apomarsupella R. M. Schust. is nested within the genus Gymnomitrion, thus Apomarsupella revoluta R. M. Schuster is treated here as Gymnomitrion revolutum (Nees) H. Philibert.



Damsholt, K. 2002. Illustrated flora of Nordic liverworts and hornworts. Nordic Bryological Society, Lund, Sweden.  Damsholt, K. 2013. The Liverworts of Greenland. Nordic Bryological Society, Lund, Sweden.  Hong, W. S. 2000: The genus Marsupella in Western North America. Lindbergia 8: 166--176.

Paton, J. A. 1999. The Liverwort Flora of the British Isles. Harley Books, Colchester, England.  Schuster, R. M. 1974. Gymnomitrion. In: R. M. Schuster. The Hepaticae and Anthocerotae of North America east of the hundredth meridian. New York, Vol. 3. Pp. 115--157.  Schuster, R. M. 1988. The Hepaticae of South Greenland. Nova Hedwigia, Beih. 92: 165--169.  Schuster, R. M. 1995. On a New Species of Gymnomitrion, G. mucrophorum Schust., sp. n. Bryologist 98: 242--245.


1. Plants usually reddish- to chestnut-brown or reddish-black; leaves seldom clearly imbricate, with marginal 1--2 cell rows usually persistent, not differentiated from intramarginal cells, not hyaline at maturity, more or less tangentially elongated (if at all), with oil-bodies; underleaves always absent.

2. Leaf margins reflexed, even in the sinuses.

3. Leaf margins uniformly, strongly widely reflexed to revolute; leaves on mature shoots longer than wide; leaf cell trigones indistinct, smaller than cell lumen, rarely confluent in rectangles ............ 1. Gymnomitrion revolutum

3. Leaf margins narrowly reflexed leaves on mature shoots usually wider than long, or as wide as long, only rarely slightly longer than wide; leaf cell trigones distinct, often as large as or even larger than cell lumen, often confluent in rectangles ....... 2. Gymnomitrion commutatum

2. Leaf margins plane, never reflexed.

4. Monoicous; plants usually small, up to 8 mm long and 0.2--0.6 mm wide; leaves appressed below and contiguous to weakly imbricate, towards the shoot apex gradually increasing in size, densely imbricate, erect to erecto-patent, not sheathing at base, not decurrent.

5. Leaves wide oval, 1.1--1.4 as long as wide; sinus rectangular to less often broadly rounded ............... 3. Gymnomitrion brevissimum

5. Leaves oblong, oblong ovate, or oblong-obovate, 1.4--1.9 as long as wide; sinus narrowly to broadly rounded .................. 4. Gymnomitrion adustum

4. Dioicous; plants medium-sized to large, 10--30 mm long and 0.3--1(--1.3) mm wide; leaves of equal size throughout, distant and patent to almost at right angles to the stem; sheathing at base, with both dorsal and ventral decurrency ......... 5. Gymnomitrion alpinum

1. Plants whitish, greenish, yellowish to brown, rarely red-brown; leaves close, frequently tightly imbricate; marginal cells in 1--3-rows usually hyaline, often variously differentiated (thin-walled and slender, or rather thick-walled), often elongated obliquely or at right angles to the margin, without oil-bodies; underleaves absent or rarely present, minute.

6. Leaves bifid (0.35--)0.4--0.55 their length ....... 6. Gymnomitrion laceratum

6. Leaves shallowly 2-lobed to 0.1--0.35 their length.

7. Leaves with mucronate lobes, ending in one elongated or 2 superposed cells (but some leaves sometimes with one lobe mucronate the other rounded), serrulate with the free apices of obliquely elongate, sharply projecting cells ..... 7. Gymnomitrion mucrophorum

7. Leaves with lobes similar, neither clearly mucronate nor serrulate; marginal cells often obliquely elongate and rectangular, but free ends projecting at most as crenulations.

8. Leaf cells in 1--2 marginal rows thin-walled and delicate, often erose.

9. Shoots greenish, yellowish to brown, at times red-brown or even reddish, terete; marginal cells of leaves elongated obliquely or at right angle to the margin, with rounded free ends protuberant to 0.3 their length .... 8. Gymnomitrion pacificum

9. Shoot white or dark grey to black, clavate to lingulate, distinctly flattened; marginal cells of leaves rectangular or rhomboid, sometimes tangentially elongated, only slightly projecting as round crenulations ..... 9. Gymnomitrion corallioides

8. Marginal leaf cells mostly thick-walled, decolorate, but normally not becoming erose with age.

10. Leaf lobes broadly rounded; underleaves sometimes present, lanceolate, shortly connate at base with one leaf ........... 10. Gymnomitrion obtusum

10. Leaf lobes obtuse to apiculate; underleaves absent .... 11. Gymnomitrion concinnatum



1. Gymnomitrion revolutum (Nees) H. Philibert, Rev. Bryol. 17(3): 34. 1890


Sarcocyphos revolutus Nees, Naturgesch. Eur. Leberm. 2: 419. 1836; Marsupella revoluta (Nees) Dumortier


Plants brownish to reddish black, forming dense patches.  Shoots 10--50 mm long, 1--2 mm wide, erect, rigid, simple or sparingly branched, rarely with the dorsally displaced branches from the dorsal part of lateral merophytes.  Rhizoids sparse.  Stems 230--300 \um in diam., in cross section with several layers of thick-walled cortical cells, the outermost with yellowish walls, the inner cells with brownish walls, grading into the collenchymatous pale medullary cells.  Leaves almost of equal size throughout, contiguous to distant, stiffly- or erect-spreading, rarely weakly imbricate, transversely inserted and oriented, sheathing basally, conspicuously decurrent ventrally and dorsally, obovate or elliptical, somewhat spatulate, 800--1050 x 650--745 \um, 2-lobed 0.25--0.40 of the length, lobes ovate-triangular, acute or obtuse at apex, sinuses acute to slightly obtuse, or rounded at the gibbous base; whole margin revolute, even in the sinuses, never erose, entire (marginal cells not protuberant).  Leaf cells of the margins 10--12 \um wide, not decolorate with age; cells of leaf lobes 12--18 x 9--11 \um, of leaf middle 15--25 x 12--14 \um, basal cells of 24--36 x 14--17 \um, the walls often somewhat thickened, with small and indistinct trigones.  Cuticle smooth to finely verruculose.  Underleaves absent.  Sexual condition dioicous.  Androecial bracts in several pairs, similar to the leaves but more concave at base.  Gynoecial bracts erect, larger than leaves, 2-lobed 0.2--0.25 of the length, obtuse, their margins not revolute.  Perianth absent.  Elaters 2-spiral, 6--8 \um in diam.  Spores yellowish brown, 12--15 \um.


Restricted to always non calcareous, damp or wet, or periodically irrigated rock faces or vertical cliff walls, in and around rock pools; low to high elevations; arctic-alpine; Greenland; B.C., N.W.T., Nunavut, Yukon; Colo.; Europe; Asia.


Gymnomitrion revolutum easily differs from all other regional representatives of the genus by the (1) rigid, rather large, mostly erect shoots, (2) brownish to reddish black pigmentation and (3) whole leaf margins including sinus broadly reflexed to revolute.



2. Gymnomitrion commutatum (Limpricht) Schiffner, Magyar Bot. Lapok 13: 304. 1915


Sarcocyphos commutatus Limpricht, Jahresber. Schles. Ges. Vaterl. Cult. 57: 314. 1880; Marsupella commutata (Limpricht) Bernet


Plants reddish- to blackish brown, forming dense patches.  Shoots 5--20 x 0.4--0.8 mm, prostrate, simple or sparingly branched.  Rhizoids sparse.  Stems 130--150 \um in diam., in cross section with 1--2 layers of equally thick-walled cortical cells of almost the same size as usually thin-walled but collenchymatous medullary cells.  Leaves usually of equal size throughout, approximate to moderately imbricate, pectinately distichous, suberect to obliquely patent, shortly decurrent dorsally, wide ovate to-rounded quadrate, usually narrow at base, concave, 470--650 x 360--650 \um, 2-lobed 0.15--0.3 of the length, with ovate-triangular obtuse lobes and erect to slightly incurved apices; sinuses acute to rarely obtuse, margins narrowly reflexed, even in the sinuses, entire (marginal cells not protuberant). Leaf cells in the middle 12--20 x 8--12 \um, with trigones large, sometimes as large as cell lumen, mostly convex, often confluent in rectangles.  Cuticle smooth to finely verruculose.  Underleaves absent.  Sexual condition dioicous. Androecia at apex of shoots, spicate, become later intercalary; gynoecia terminal on main shoots. Androecial bracts in several pairs, imbricate, similar to the leaves but strongly concave, slightly reflexed at margins. Gynoecial bracts erect, similar to leaves but larger, 740–780 x 700–780 \um, concave, margins of bracts and sinus reflexed. Perianth absent.  Elaters 2-spiral.  Spores light brown, minutely verruculose, 10--17 \um. 


On thin soil over granite rocks and in the niches between boulders on rock fields, or on bare damp soil in fell-fields, late snow areas and along streams; low to high elevations; arctic-alpine; Greenland; B.C., Yukon; Alaska, Wash.; Europe; Asia.


Gymnomitrion commutatum is most closely related in appearance to G. alpinum and G. brevissimum, but easily differs by the leaf margins narrowly reflexed for all the length, including sinus.


3. Gymnomitrion brevissimum (Dumortier) Warnstorf, Hedwigia 53(3): 196. 1913


Acolea brevissima Dumortier, Syll. Jungerm. Europ.: 76. 1831; Marsupella brevissima (Dumortier) Grolle


Plants reddish brown or sometimes almost fuscous black, forming dense patches.  Shoots erect or strongly ascending, subterete, slender and small-leaved below, 0.15 mm wide below and becoming 4--10 x 0.45--0.7 mm, often irregularly branched, rarely simple.  Rhizoids sparse, except on older parts of plant, hyaline or reddish.  Stem lacking thick-walled cortical layer.  Leaves small, appressed and contiguous to weakly imbricate below, gradually increasing in size and becoming distinctly imbricate towards the shoot apex, sometimes antically secund, erect, erecto-patent to erect-imbricate, not tightly appressed, oval to oval-oblong to oblong-elliptical, not or only slightly decurrent, concave, 2-lobed for 0.2--0.35 of the length; sinuses acute to rectangular, lobes ovate-triangular, acute or subacute at apex, often slightly incurved; the upper leaves of sterile shoots 250--550 x 220--450 \um; the leaf margins not reflexed, entire (marginal cells not protuberant), normally not decolorate, never erose.  Leaf cells at the margins 9--12 \um wide, not differentiated from interior cells; median cells 10--20 x 10--17 \um; cell walls somewhat thickened, with large, distinct to bulging trigones.  Cuticle smooth.  Underleaves absent.  Sexual condition paroicous or autoicous, with pure male and female plants possibly occurring.  Androecial bracts in 2--3 pairs, below the gynoecia or on separate branches, rounded-ovate to subrotundate, often armed with an obtuse lobe-like tooth towards base of dorsal margin.  Gynoecial bracts similar to leaves but much wider, more concave and less deeply divided than leaves, ca. 0.15 2-lobed, with apiculate lobes and acute to lunate sinuses.  Perianth absent or partly present.  Elaters 8--10 \um wide, 3--4-spiral.  Spores pale reddish brown, almost smooth or finely papillate or granulate, 8--15 \um.


On bare damp soil in late snow areas and along streams, often on rocky ground, apparently decidedly oxylophytic, absent in calcareous regions; low to high elevations; arctic-alpine and subalpine; Greenland; Alta, B.C.; Alaska, Wash.; Europe; Asia.


4. Gymnomitrion adustum Nees, Naturgesch. Eur. Leberm. 1: 120. 1833


Marsupella adusta (Nees) Spruce


Plants green, olive green, dark greyish, brownish green, dark brown or reddish brown, forming dense patches.  Shoots 2--6 x 0.25--0.5 mm, simple or irregularly branched, strongly to weakly compressed, erect or ascending. Rhizoids hyaline or reddish or purplish, numerous in older parts, sparse in upper parts of branches.  Leaves gradually increasing in size towards apex, almost erect or suberect or rarely erecto-patent, imbricate yet loosely appressed, weakly canaliculate, with base somewhat sheathing the stem, with the slightly decurrent margins, elliptical or less often ovate, longer than wide, 350--500 x 200--320 \um, 2-lobed 0.16--0.3 of the length; sinus narrowly to broadly rounded or sometimes acute, lobes triangular to ovate, with apex sometimes incurved, narrowly to broadly rounded, or obtuse, or sometimes one or both lobes acute with a single apical cell.  Leaf cells at the margins 9--12 \um wide, not differentiated from interior cells; median cells 20--25 x 12--15 \um; walls somewhat thickened, with distinct to large bulging trigones, especially in leaf lobes.  Cuticle smooth or weakly papillose.  Underleaves absent.  Sexual condition paroicous, or rarely synoicous, only known fertile.  Androecial bracts in 2--3 pairs, below the gynoecia, similar to the leaves, but larger, somewhat ventricose at base.  Gynoecial bracts up to 0.7mm long, triangular-rotundate, as wide as or slightly wider than long, widest at base, concave, 2-lobed 0.15--0.25 of the length, with obtuse lobes; sinuses obtuse to broadly rounded.  Perianth absent.  Elaters 6--9 \um wide, 4-spiral.  Spores reddish brown, almost smooth, 7--12 \um.


On acidic to mildly base-rich stones or rock walls in or beside flushes, small streams and lakes, on mountain screes, sometimes on soil on ledges, in crevices and on steeply sloping wet rocks on montane crags, on stones and soil irrigated by water from late snow beds; mostly low to moderate elevations; subalpine and alpine; B.C.; Oreg.; Europe; Asia.


The reports of Gymnomitrion adustum from eastern and western Canada were rejected by J. Váňa et al. (2010: 17); however, the specimens from Oregon (MO) and British Columbia (UBC) here studied cannot be attributed for any other known species. From the most closely related species G. brevissimum these specimens differ in morphology and molecular features.


5. Gymnomitrion alpinum (Gottsche ex Husnot) Schiffner, Osterr. Bot. Z. 53: 280. 1903


Sarcocyphos alpinus Gottsche ex Husnot, Hepaticol. Gall. 1: 13. 1875; Marsupella alpina (Gottsche ex Husnot) Bernet


Plants glossy, greenish or reddish brown to dark brown, copper red or almost black, sometimes pale yellowish or reddish green, forming loose to rather dense mats or carpet- to cushion-like tufts.  Shoots horizontal and pendent to suberect or erect, 10--40 x 0.3--1.3 mm, compressed and pectinate or sometimes subterete and arcuate or filiform, simple or sparingly branched. Rhizoids sparse, colorless to purplish.  Stem in cross section with one layer of brownish thick-walled cortical cells of almost the same size as the medullary cells.  Leaves of equal size throughout, distant and patent to almost at right angles to the stem, rarely slightly imbricate, rounded elliptical or ovate, or subquadrate, with erect concave base sheathing the stem, with both dorsal and ventral decurrency, transversely to somewhat obliquely inserted or sometimes horizontal and nearly plane, up to 0.8 mm long, nearly as wide as to slightly wider than long; margins sometimes eroded and becoming denticulate with age; 2-lobed 0.2--0.4 of the length; sinuses acute to obtuse or broadly rounded, sometimes rectangular, lobes ovate-triangular, broadly to narrowly rounded or obtuse or sometimes subacute, often slightly incurved at apex.  Leaf cells at margins 8--10 \um wide, almost not differentiated from interior cells; median cells 10--19 x 8--16 \um, with large trigones often confluent in rectangles.  Cuticle smooth.  Underleaves absent.  Sexual condition dioicous, with male plants often in separate tufts.  Androecial bracts in several pairs, terminal or intercalary on main shoot, not or slightly larger than adjacent leaves, imbricate or occasionally distant, concave, ovate to oblong, ca. 0.4--1.0 mm wide, 2-lobed up to 0.5 the length, lobes usually rounded.  Gynoecial bracts larger than the leaves, concave, rounded-triangular, 2-lobed 0.15--0.2 of the length, with obtuse to rounded lobes.  Perianth slender, inflated, plicate.  Elaters 7--8 \um wide, 2--4-spiral.  Spores almost smooth, 10--11 \um.


On wet or periodically irrigated or moist to rather dry acidic to mildly base-rich rocks, on steeply sloping to vertical or nearly flat surfaces on mountain crags, boulders and rock slabs, also on small rocks and gravelly soil irrigated by water from late-lying snow; low to high elevations; subalpine and alpine; B.C.; Alaska, Wash.; Europe; Asia.


The specimens of Gymnomitrion alpinum from Alaska, British Columbia and Washington (MO and UBC) differ from European specimens of G. alpinum by (1) smaller rigid shoots, (2) deep copper red to almost vinaceous pigmentation, (3) pectinately distichous vegetative leaves which are almost rectangular and thick (with cell lumen higher than wide in cross section), often with a very narrow hyaline border of strongly thickened and decolorate external walls of marginal cells, also by (4) rather small median leaf cells, mostly 8--13 \um wide, and (5) perianth mouth crenulate with highly projecting rounded free ends of marginal cells and 1-celled teeth of thin-walled, somewhat elongated marginal cells which are free for all the length. These Western North American specimens perhaps represent a separate subspecies of G. alpinum, rather than a morphological extreme. This question needs further investigation.


6. Gymnomitrion laceratum (Stephani) Horikawa, Acta Phytotax. Geobot. 13: 212. 1943


Acolea lacerata Stephani, Sp. Hepat. 6: 78. 1917


Plants bright gray-green to glaucous, yellowish or weakly brownish tinged, forming dense tufts. Shoots flexuous and rigid, julaceous to filiform, 3--10 x 0.2--0.5 mm, decumbent, with distal portions scarcely ascending, simple or irregularly branched. Rhizoids frequent, hyaline, extremely slender and long, extending to near the stem apex. Stems 108--150 \um in diam., with the cortical cells thin-walled like medullary cells and subequal to them in size (in cross section).  Leaves densely imbricate, on occasional suberect shoots virtually transverse, on decumbent shoots slightly but distinctly succubous, narrowly ovate to oblong-ovate, ca. 360--450 x 270--360 \um, entire-margined, 2-lobed 0.35--0.55 their length; sinuses acute, the notch usually quite narrow, lobes to 1.2--1.5 as wide as long, narrowly triangular to ovate-triangular, mostly acute, terminated at apex by a single, slightly to moderately elongated cell, or by 2 cells in a row, rarely blunt and terminated by 2 juxtaposed cells.  Leaf cells at the margins equally thick-walled (with walls up to 0.5 the cell width) and hyaline, in 2 or more rows, 16--24 x 12--15 \um, forming a conspicuous border distinct from the interior cells; cells in the middle 17--22 x 15--17 \um, with trigones small or almost absent, cells in the base 26--38 x 22--24 \um. Cuticle minutely verruculose.  Underleaves usually absent, rarely present, minute (2--8 cells in size), lanceolate to triangular, at times slightly or rather strongly connate with the postical base of the leaf on one side of stem.  Sexual condition dioicous. Gynoecial and androecial shoots both more clavate than vegetative shoots. Androecial bracts in several pairs, concave, 2-lobed 0.4--0.45 their length, broader than leaves. Gynoecial bracts deeply lacerate-laciniate, divided into a series of laciniae ending in slender cilia, uniseriate for a length of 4--9 cells. Perianth absent.  Elaters 2-spiraled, ca. 6--7 \um in diam. Spores brown, verruculose, 13--15 \um in diam.


In shaded crevices of vertical or steeply sloping, damp rocks, associated mostly with acidophytic hepatics; low to high elevations; Tenn.; Mexico; Asia.


Gymnomitrion laceratum easily differs from all other regional representatives of the genus by the bright pigmentation, abundant long rhizoids and deeply 2-lobed leaves.



7. Gymnomitrion mucrophorum R. M. Schuster, Bryologist 98: 243. 1995


Plants light green, whitish when dry, with yellowish or reddish brown pigmentation, often nitid.  Shoots terete, threadlike, simple or irregularly branched. Leaves oblong to oblong-ovate, 605--675 x 480--540 \um, widest in basal 0.2, 2-lobed 0.15--0.28 of the length, with a sharply acute sinus, with variable lobes; one lobe always sharply acute, the other sometimes irregularly truncate or blunt or emarginate; sharp lobes ending in a single, thick-walled elongate cell (30--38 x 11--15 \um, ca. 2--3:1) or two superposed cells, below the tips often with 2-several erect or suberect cells with thick-walled, tapered free apices. Lobe margins plane, serrulate. Leaves without defined, narrow border, but with distal half and margins decolorate at maturity, formed of suberect to obliquely somewhat elongate thick-walled cells with bulging trigones, with strongly thick-walled free apices obtusely to angularly projecting.  Leaf cells at the serrulate margins 25--32 x 12--15 \um, within the thick-walled border strongly collenchymatous, subisodiametric, with bulging or confluent trigones, 17--22 x 15--19 \um; median cells rather thin-walled, with nodulose trigones, 23--30 x 20--25 \um.  Cuticle asperulate to verrucate or vermiculate.  Underleaves absent.  Other features unknown.


On silty soil over boulders in a humid boulder field in late snow area; moderate elevations; subalpine; Greenland; Alaska.


Gymnomitrion mucrophorum differs from the most closely related species G. concinnatum mainly by the serrulate leaf apices. The distinction of G. mucrophorum may need to be reinvestigated using molecular methods when more gatherings from the type locality have accumulated.



8. Gymnomitrion pacificum Grolle, Trans. Brit. Bryol. Soc. 5: 92, fig. 2f--k. 1966


Plants greenish, yellowish or reddish brown, rigid, ca. 10--20 mm long, usually in compact patches.  Shoots about 0.25--0.7 mm in diam., blunt or rounded at tips, smooth through erect-appressed leaves, but with leaf margins not difficultly discernible (glistening when dry), filiform to julaceous, almost terete, simple or sparingly branched.  Stems 110--120 \um in diam., with the cortical cells little thickened.  Leaves dense, almost transverse, very closely, regularly inserted, not dorsally secund to barely so, closely imbricate and erect-appressed, canaliculate, broadly triangular-ovate, sometimes wider than long, amplexicaul, narrowed to the base, ca. 410--580 x 260--600 \um, 2-lobed for 0.09--0.25 their length; sinuses rectangular to acute, lobes broadly triangular; margins slightly to distinctly crenulate with a partially erose border of 1--2(--3) rows of hyaline cells oriented with their long axes at an oblique angle with leaf margins, with rounded free ends protuberant for 0.3 of the length of the cells; leaf apices obtuse to almost rounded through erose margins, rarely apiculate (ending by an acute cell).  Leaf cells at margins 14--21 x 8--13 \um, mostly thin-walled; submarginal cells with light fuscous pigmentation, with bulging trigones and thickened intermediate walls, or almost equally thick-walled, in lobes 15--21 x 11--16 \um, at base 14--29 x 11--19 \um.  Cuticle minutely verruculose.  Underleaves absent.  Sexual condition dioicous, with clavate gynoecia and androecia.  Androecial bracts in numerous pairs, slightly ventricose, ovate, 350--400 x 320--450 \um.  Gynoecial bracts much larger the leaves, widely ovate, strongly convolute.  Perianth absent.  Elaters 7--10 \um wide, 2--3-spiral.  Spores brownish, almost smooth, 10--14 \um.


On moist rocks and rocky slopes; low and moderate elevations; arctic-alpine; B.C., Alaska; Asia.


Dry plants of Gymnomitrion pacificum easily differ from all other regional species of the genus by the rigid, almost terete, filiform to julaceous shoots with yellowish or reddish brown pigmentation and strongly appressed leaves with glistening hyaline margins consisting of mostly thin-walled cells.



9. Gymnomitrion corallioides Nees, Naturgesch. Eur. Leberm. 1: 118. 1833


Plants silvery or whitish-grey, yellowish brown or pale to dark grey or nearly black, forming loose to dense turfs or mats.  Shoots 10--20 x 0.3--0.7 mm, simple to frequently branched, below with stolons, clavate to linear and dorsiventrally compressed, rarely filiform and almost terete, erect to prostrate, frequently branched.  Rhizoids whitish, virtually lacking or sparse above, numerous below.  Leaves densely imbricate, erect and appressed, 540--700 x 580--660 \um, suborbicular or broadly to rotund ovate, widest in basal third, concave, margins edentate or weakly crenulate, 2-lobed 0.1--0.15 of the length; sinuses obsolete, acute to rectangular or rounded, lobes scarcely differentiated, short and broad, rounded or subacute, with a sharp projecting apical cell, soon ill-defined through destruction of apical and marginal cells. Margins never revolute but soon becoming decolorate, marginal 2--3 rows of hyaline cells often eroded.  Leaf cells at the margins 12--15 \um wide, in 1--3 rows rectangular to isodiametric or sometimes tangentially elongate, thin-walled, slightly projecting as crenulations; submarginal cells in several rows hyaline, thick-walled, with large, often confluent trigones, especially in the distal part of the leaf, 14--19 \um wide; median cells 18--26 x 16--23 \um; walls slightly thickened, with distinct trigones.  Cuticle smooth or minutely papillose.  Underleaves absent.  Sexual condition dioicous, with strongly clavate, not or hardly dorsiventrally compressed female shoots.  Androecial bracts becoming intercalary in several pairs, similar to leaves, but less closely imbricate, with reflexed apices, often with an obtuse tooth at the dorsal margin.  Gynoecial bracts in several pairs; outer bracts dentate, with reflexed margin, with acute sinuses and acute lobes crenulate to dentate-ciliate at margins, with the marginal cells often radially elongate; inner bract irregularly lobed and dentate.  Perianth absent.  Elaters 7--8 \um wide, 2-spiral.  Spores reddish brown, verruculose, 12--16 \um.


Near mountain summits, rarely on rocky slopes in mountain forests; on exposed, mostly acidic or base-rich windswept cliffs and rock walls, on a thin layer of soil or humus on moist or wet boulders, on ledges, on ridges and in crevices, also on stones and stony soil, on bare exposed heath soil, between rocks in fell-fields, often closely adjacent to late-lying snow or permanent ice caps; low to high elevations; arctic-alpine; Greenland; B.C., Nfld. and Labrador, N.W.T., Nunavut, Yukon; Alaska, Colo., Maine, N.H.; Europe; Asia.



10. Gymnomitrion obtusum Lindberg, Morgonbladet (Helsinki) 1877(30): 2. 1877


Plants whitish or pale greyish green, greyish brown, yellowish or reddish brown on distal exposed portions of shoots, forming compact cushion-like turfs or dense mats.  Shoots erect to prostrate, 10--20 x 0.2--0.6 mm, linear and almost terete, occasionally filiform, simple or with few branches.  Rhizoids scarce, only occurring near base and at stolons.  Leaves densely imbricate, erect and symmetrically appressed, subtransversely inserted and oriented, broadly elliptical or obcordate, strongly concave, 550--900 x 550--775 \um, 2-lobed 0.16--0.25 their length; sinuses generally sharp, lobes evenly rotundate, sometimes narrowly rounded, rarely subacute and minutely apiculate, margins nearly always distinctly and rather evenly crenulate, almost to the base, marginal 1--2 rows of cells and distal portions of leaves soon becoming hyaline, with reflexed dorsal margin.  Leaf cells at the margins normally persistent, quadrate-rectangular, thick-walled, sometimes slightly elongated obliquely or at right angle to margin, 12--15 \um wide; submarginal cells almost uniformly thick-walled; median cells 18--27 x 14--20 \um, thin-walled, with distinct to rather large bulging trigones, in lobes often confluent in rectangles. Cuticle minutely and densely verruculose.  Underleaves absent or rarely present as lanceolate vestiges, shortly connate at base with one leaf.  Sexual condition dioicous; often fertile; male and female shoots somewhat clavate and slightly compressed.  Androecial bracts terminal or intercalary, in several pairs, with rounded lobes; similar to the leaves, but larger; saccate, closely imbricate, with reflexed apices and dorsal margin and sometimes margins of sinus.  Gynoecial bracts in 3--4 pairs; outer bracts larger than the leaves, irregularly and shallowly lobed, with rotundate or sometimes subacute lobes, with reflexed crenulate margins; inner bracts more delicate, smaller than leaves, with truncate, acute or shortly ciliate lobes and crenulate-dentate margins.  Perianth absent.  Elaters 7--11 \um wide, 2-spiraled.  Spores reddish brown, minutely verruculose, 12--16 \um.


On periodically moist, acidic to base-rich rock walls in ravines, on steeply sloping rock outcrops and crags, on ledges and stones, usually on the surface or on thin soil over, on exposed dry rocks in humid habitats, in the vicinity of late-lying snow and on mountain summits, on boulders in and beside rivers; low to moderate elevations; subalpine and alpine; Greenland; B.C.; Alaska, Mont., Oreg., Wash.; Europe.


Gymnomitrion obtusum is easily distinguished from all other regional species of the genus by the broadly rounded apices of leaf lobes with crenulate, partly reflexed margins of thick-walled hyaline cells.



11. Gymnomitrion concinnatum (Lightfoot) Corda, Deutschl. Fl., Abt. II, Cryptog. 19: 23. 1830

Jungermannia concinnata Lightfoot, Fl. Scot. 2: 786. 1777, nom. conserv.


Plants pale or whitish green or yellow-green, with distal portions of leaves more often yellowish brown to dark or chestnut brown (except for decolorate, whitish leaf tips and narrow margins), forming loose to dense, sometimes cushion-like turfs or thin mats.  Shoots 5--20 mm long, 0.25--0.7 mm wide, linear or weakly clavate and dorsiventrally compressed, rarely filiform and almost terete, erect to prostrate, simple or irregularly branched.  Rhizoids whitish, sparse, present below and from stolons.  Leaves erect or suberect, slightly appressed, imbricate, barely dorsally secund, suborbicular to broadly elliptical or ovate, 400--1000 x 350--700 \um, concave, 2-lobed 0.1--0.35 of the length; sinuses acute, rarely rounded, or moderately wide to rectangular, lobes triangular or ovate-triangular, subacute or acute, or rarely somewhat obtuse, or weakly apiculate, terminated of 1--2 superposed cells, with one or both lobes sometimes rounded; margins edentate or crenulate to subserrate, leaf apex and marginal 1--3 rows of cells soon becoming hyaline, but normally not erose with age.  Leaf cells at the margins thick-walled, isodiametric, or sometimes elongated tangentially or at right angle to the margin, 12--15 \um wide; median cells 20--28 x 15--24 \um; walls slightly thickened, with bulging, often almost confluent trigones.  Cuticle smooth or minutely papillose.  Underleaves absent.  Sexual condition dioicous, with clavate male and female shoots.  Androecial bracts becoming intercalary, in several pairs, similar to leaves, but larger, less closely imbricate, saccate at base, with lateral margins narrowly recurved.  Gynoecial bracts in several pairs, outer bracts larger than the leaves and more deeply divided; inner bracts with acute or ciliate lobes, with margins irregularly dentate-ciliate and weakly crenulate.  Perianth absent.  Elaters 7--8 \um wide, 2-spiral.  Spores reddish brown, verruculose, 12--15 \um in diam.


Typically in exposed, insolated sites on mountain summits and slopes; on rather dry to wet, acidic to base-rich cliffs and crags, on ledges and in crevices, on boulders, on gravelly, sandy or peaty soils, often on thin layer of soil or humus, on screes and areas of late-lying snow; low to high elevations; arctic-alpine; Greenland; Alta, B.C., Nfld. and Labrador, N.W.T., Nunavut, Que., Yukon; Alaska, Colo., Maine, Mont., N.H., Oreg., Wash.; Europe; Asia.


The specimens of Gymnomitrion concinnatum from New Hampshire (in MO) differ from typical plants by (1) slender plants, (2) olive to golden brown pigmentation, (3) leaf apices partly recurved in dry plants, (4) weakly differentiated marginal cells of leaf lobes with slightly thickened walls and rather large size, 13--18 \um wide, and (5) more large size of median leaf cells, up to 27 \um wide. These specimens perhaps represent a leptodermous modification of G. concinnatum, but this question needs further investigation when more gatherings have accumulated.


OTHER REFERENCES  Crandall-Stotler, B., R. E. Stotler and D. G. Long. 2009. Phylogeny and classification of the Marchantiophyta. Edinb. J. Bot. 66: 155--198.  Crandall-Stotler, B. and R. E. Stotler. 2017. A Synopsis of the Liverwort Flora of North America North of Mexico. Annals of the Missouri Botanical Garden 102(4): 574--709.  Kitagawa, N. 1963. A revision of the family Marsupellaceae of Japan. Journal of the Hattori Botanical Laboratory 26: 76--118.  Mamontov, Yu.S., N.A. Konstantinova, A.A. Vilnet, A.D. Potemkin, E.V. Sofronova and N.S. Gamova. 2018. On resurrection of Marsupella parvitexta Steph. (Gymnomitriaceae, Marchantiophyta) as a semi cryptic species of the genus Gymnomitrion. Nova Hedwigia 106(1-2): 81–101.  Potemkin, A.D., Yu.S. Mamontov and N.S. Gamova. 2017. Gymnomitrion fissum (Gymnomitriaceae, Marchantiophyta) – a new species with fissured leaf surface from China. Novosti Sist. Nizsh. Rast. 51: 274–280.  Schumacher, R. and J. Váňa. 2005. Identification keys to the liverworts and hornworts of Europe and Macaronesia (distribution and status). Poznań, Poland.  Schuster, R. M. 2002. Austral Hepaticae Part II. Nova Hedwigia, Beih. 119: 562--577.  Shaw, B., B. Crandall-Stotler, J. Váňa, R. E. Stotler, M. von Konrat, J. J. Engel, E. C. Davis, D. G. Long, P. Sova and A. J. Shaw. 2015. Phylogenetic Relationships and Morphological Evolution in a Major Clade of Leafy Liverworts (Phylum Marchantiophyta, Order Jungermanniales): Suborder Jungermanniineae. Systematic Botany 40(1): 27--45.  Söderström, L., A. Hagborg, M. von Konrat et al. 2016: World checklist of hornworts and liverworts. PhytoKeys 59: 1--828. 



2. MARSUPELLA Dumortier, Commentat. Bot., 114. 1822 * [Latin marsupium alluding to the brood pouch]


Nadezhda A. Konstantinova


Plants small to medium size, rarely large, often growing erect, green or more often with tints of red and red-brown to blackish secondary pigmentation. Stems simple or with a few lateral intercalary branches, but often apparently ventral intercalary, often with numerous stolons, in cross section with hyalodermis or this absent. Rhizoids sparse, near base and on stolones. Leaves transversely inserted, occasionally succubous, in two rows, 2-lobed for less than 0.5(--0.6) of their length, lobes of equal or almost equal size, entire. Laminal cells distinctly collenchymatous from very small, only 7--8 /um wide, to relatively large (20--30 /um) with 2--3(--4) relatively large oil bodies per cell. Underleaves absent. Sexual condition monoicous or dioicous. Androecium terminal or intercalary, male bracts larger than the leaves and more concave; antheridial stalk 1--2-seriate. Gynoecium terminal at main shoot, Female bracts much larger and less deeply divided than leaves, in 2--3 pairs, bracteole absent. Perigynium well developed. Perianth small, shortly immersed, perianth mouth crenulate. Capsule spheric-ovoid; valves 2--3-stratose; inner layer with radial or semiannular thickenings, epidermal layer with radial nodular thickenings. Spores small, 7--13 /um. Elaters 2--4-spiral.


Species 33 (15 in the flora): bipolar, widely distributed in Arctic and mountainous areas, North America, Europe, Asia, Pacific Ocean Islands (Hawaii), subantarctic islands.



Hong, W. S. 1982. The genus Marsupella in western North America. Lindbergia 8 (3): 166--176.

Bakalin, V. A., Fedosov V. E., Fedorova A. V. & Nguyen V. S. 2019.  Integrative taxonomic revision of Marsupella (Gymnomitriaceae, Hepaticae) reveals neglected diversity in Pacific Asia. Cryptogamie,Bryologie 40 (7): 59-85.

Mamontov, Yu. S., A.A. Vilnet, N.A. Konstantinova & V. A. Bakalin 2019. Two new species of Gymnomitriaceae (Marchantiophyta) in the North Pacific // Botanica Pacifica. A journalplant science and conservation. V. 8(1):67--80 DOI: 10.17581/bp.2019.08113

Schuster, R. M. 1974. The Hepaticae and Anthocerotae of North America east of the hundredth meridian. New York. Vol. 3: 1--880.

Schuster, R. M. 1974. The Hepaticae of West Greenland from ca. 66ºN to 72ºN Meddelelser om Grønland udgivene af Kommissionen for videnskabelige undersøgelser i Grønland. Bd. 199. Nr.1 370 p.

Schuster, R. M. 1988. The Hepaticae of South Greenland. Beih. Nova Hedwigia. Heft 92. 255 pp.

Söderström, L., A. Hagborg, M. von Konrat, S. Bartholomew-Began, D. Bell, L. Briscoe, E. Brown, D. C. Cargill, D. P. Costa, B. J. Crandall-Stotler, E. D. Cooper, G. Dauphin, J. J. Engel, K. Feldberg, D. Glenny, S. R. Gradstein, X. He, J. Heinrichs, J. Hentschel, A. L. Ilkiu-Borges, T. Katagiri, N. A. Konstantinova, J. Larraín, D. G. Long, M. Nebel, T. Pócs, F. Puche, E. Reiner-Drehwald, M. A. M. Renner, A. Sass-Gyarmati, A. Schäfer-Verwimp, J. G. S. Moragues, R. E. Stotler, P. Sukkharak, B. M. Thiers, J. Uribe, J. Váňa, J. C. Villarreal, M. Wigginton, L. Zhang  and R.-L. Zhu. 2016. World checklist of hornworts and liverworts. PhytoKeys 59: 1--821.

Stotler, R. E. & B. J. Crandall-Stotler. 2017. Synopsis of the Liverwort Flora of North America North of Mexico. Ann. Missouri Bot. Gard. 102: 574--709.

Váňa, J., L. Söderström, A. Hagborg, M. von Konrat & J. J. Engel. 2010. Early land plants today: Taxonomy, systematics and nomenclature of Gymnomitriaceae. Phytotaxa 11: 1--80.

Vilnet, A. A., N. A. Konstantinova and A. V. Troitsky. 2007. On molecular phylogeny of Gymnomitriaceae H. Klinggr. (Hepaticae). Computational Phylogenetics and Molecular Systematics. CPMS” 2007, Conference proceedings. KMK, Moscow, pp. 24--26.


1. Plants densely leaved, almost worm-like in appearance, with appressed and closely imbricate leaves; marginal leaf cells decolorate in 1--2 rows, often erose; leaf lobes abruptly apiculate, often ending in two superposed cells or only one but then elongated cell …..……………….…… ..2. Marsupella apiculata

1. Plants with more or less remote leaves, if imbricate often not hiding the stem; marginal cells not decolorate or decolorate only near tips, not erose; leaf lobes obtuse to rarely acute, not abruptly apiculate, ending in one cell, rarely in two cells but then without decolorate margin.

2. Leaves hemispheric to spoon-shaped, almost unlobed or just shallowly retuse ………………………………………………………………………3. Marsupella arctica

2. Leaves concave or canaliculated, not hemispheric or spoon-shaped, distinctly divided into two lobes

3. Leaves with usually lunate sinus, rarely rotundate; small plants with leafy shoots not exceeding 0.5 mm wide……………. ………………………….7. Marsupella condensata

3. Leaves with acute or rounded sinus; plants medium sized or if small then always with acute sinus. 

4. Plants julaceous with leaves on sterile shoots not or slightly broader than stem, Cephaloziella-like.

5. Leaves appressed, more or less canaliculate, longer than wide, margins of male and Female bracts crenulate or dentate …..….. 15. Marsupella stableri

5. Leaves patent to suberect and erect, concave, with often incurved lobes, usually wider than long, male and Female bracts edentate ………………. 5. Marsupella boeckii

4. Plants with leaves much broader than stem, not Cephaloziella-like.

6. Paroicous or autoicous, usually fertile.

7. Plants green to somewhat brownish, never blackish or black;  leaves unequally two-lobed, with reflexed margins ……………………………………………. 10. Marsupella paroica

7. Plants often blackish or black; leaves equally two-lobed with plane margins.

8. Plants usually more than 6 mm high and 600 /um wide, deep or blackish green and rather scorched; leaves spreading, sometimes squarrose, with sheathing base and often purplish red rhizoids ……………………11. Marsupella sparsifolia

8. Plants very small with sterile shoots not exceeding 5 mm high and 450 /um wide, not scorched; leaves lacking sheathing base, rhizoids colorless.

9. Leaf lobes acuminate, divergent, often ending in apices of one elongated or 2--3 superposed cells; marginal leaf cells small, 9--11 /um clearly different from innermost cells of leaves; vegetative shoots with flattened apices ………… 13. Marsupella spiniloba

9. Leaf lobes not divergent, ending in one short cell, rarely 2 superposed cells; marginal cells not ot slightly different from the innermost; apices of vegetative shoots not flattened ……………………………… 14. Marsupella sprucei

6. Dioicous or not fertile.

10. Leaf margins more or less distinctly narrowly recurved on one or both sides.

11. Leaves wider than long, divided 0.1--0.2 of their length, lobes rotundate to obtuse, leaves bistratose near base…  ……….. ………………………….3. Marsupella aquatica

11. Leaves as wide as long, divided 0.2--0.3 of their length, lobes broadly triangular, obtuse or obtusely pointed, leaves unistratose near base…………….. 8. Marsupella emarginata

10. Leaf margins plane.

12. Plants deep or blackish green, without any trace of red pigmentation, often appears scorched; leaves sheating the stem, stem with hyalodermis 12. Marsupella sphacelata

12. Plants never appearing scorched, leaves not sheating the stem, stem without distinct hyalodermis,.

13. Leaf sinus normally not deeper than 0.25 of leaf length, some leaves with lobe-like tooth at the base of ventral side ………………….1. Marsupella aleutica

13. Leaf sinus 0.25--0.45 of leaf length, lobe-like tooth at the base of ventral side absent

14. Leaves rounded-quadrate, sinus acute, lobes triangular, acute ... 9. Marsupella funckii

14. Leaves wider than long, sinus obtuse, leaf lobes narrowly ovate-triangular, obtuse to rounded . …….. …….6. Marsupella bolanderi


1. Marsupella aleutica Mamontov, Vilnet, Konstant. & Bakalin, Bot. Pacif. 8(1):71, figs. B, D. 2019


Plants olive green to yellow brownish, shoots up to 5 mm long and 500--600 to 700 /um in fertile shoots. Stem 125 μm in diameter, in cross section with two layers of hyaline, thin- or thick-walled and slightly tangentially elongated cells 15 x 17--22(--25) μm, only slightly smaller and thin-walled innermost cells and cells of the medulla. Rhizoids few. Leaves suborbicular to often wider than long transversely or somewhat obliquely inserted, slightly to strongly imbricate conduplicate to canaliculate; sinuse to 0.2--0.25 of leaf length, acute to rounded sometimes slightly narrowly reflexed at base. Lobes slightly unequal broadly triangular mostly acute, ventral part of leaves with rounded bases, often with one to several leaves per shoot with a lobe-like tooth.  Cells of leaf margin (8--)10--13/um, in the middle 15--17(--20) x (10--)13--15(--17) /um. Oil bodies not known. Sexual condition dioicous. Female bracts from almost subquadrate  to 1.4 as wide as long, bifid to 0.15--0.2 with acute and slightly narrowly reflexed at base sinuses. Perigynium low in unfertilized plants, juvenile perianth much shorter than bracts. Otherwise unknown.

Alpine, known only from type locality in Simeonof Island (Shumagin Islands) where grow in “at the higher elevations, where the crowberry heath forms continuous slopes”; Alaska.

Marsupella aleutica (Mamontov et al. 2019) is a recently described and poorly known species. It can be confused with M. funckii.  It differs from the latter in shorter sinuses that are 0.2--0.25 of leaf length vs. 0.33--0.45 in M. funckii, smaller cells of margins, uneven lobes of leaves vs. equal sized lobes of leaves in M. funckii, and presence at least in some leaves of a lobe-like tooth that never occus in M. funckii.


2. Marsupella apiculata Schiffner, Osterr. Bot. Z. 53(6): 249, pl. 4, figs. 8--16. 1903

Gymnomitrion apiculatum (Schiffner) Müller Frib., Hedwigia 81: 113. 1942


Plants brownish green in distal parts, usually fuscous-brown to reddish tinged, to dull or brick red, 10--15(--20) x (0.33--)0.38--0.55 mm in fertile shoots, to 750 /um wide when densely leaved, such that individual leaves are hardly discernible. Stem soft-textured, ca. 175--210 /um with cortical cells hardly thick-walled; sterile shoots subequally wide, never strongly clavate, near the tip often narrowed and slightly pointed. Rhizoids few, mainly at the base of shoots. Leaves densely imbricate and erect-appressed, distinctly 2-ranked, hardly discernible, broadly cordate-ovate and cordate-subquadrate, wider than long, 420--575 x 440--650(--720) /um.  Sinus acute, rectangular, obtuse-angular, rarely lunulate (0.1--)0.2  of the leaf length.  Lobes sharply apiculate, apiculus composed of 1--2 superposed often elongated hyaline cells. Leaf cells along the margin decolorate and hyaline, often eroded, 10--15 /um; in the middle 16--19(--21) x 18--24(--30) /um, with scarcely to distinctly bulging trigones; cuticle smooth. Oil-bodies (1) 2(3--4) per cell, faintly granular, mostly 9--12 (--14) x 5--7 /um. Sexual condition dioicous. Male branches similar to the sterile, but slightly larger and subclavate, bracts slightly larger than leaves and more concave at base, bracts in several pairs, antheridia 2--3 per bract. Female branches abruptly clavate, the bracts and subfloral leaves much longer than sterile leaves, margins of bracts broadly hyaline near the base, crenulate to denticulate. Perianth plicate distally, low, equal in height to subtending Perigynium, with denticulate mouth. Capsule spheric, 2-stratose, both layers with nodular thickenings; elaters 7--8 /um, 2-spiral. Spores almost smooth, reddish brown, to 10 /um. 

Arctic-montane, mostly oceanic and suboceanic, restricted to non-calcareous sites continuously irrigated by water from melting snow; Greenland; B.C., Yukon; Alaska; Europe; Asia (South Siberia, Far East of Russia, Japan); Atlantic Islands (Iceland, Jan Mayen, Svalbard).


Marsupella apiculata can be confused with Gymnomitrion concinnatum or Marsupella condensata. From Gymnomitrion concinnatum it differs in (1) often reddish tinged shoots that never occur in G. concinnatum; (2) often erose margins vs. entire margins in G. concinnatum; (3) smooth cuticle vs. finely verruculose in G. concinnatum; (4) more obtuse sinus vs. acute deeper than 0.2 leaf length in G. concinnatum; and (5) presence of well-developed perianth vs. absent in G. conncinatum. For differences from M. condensata, see that species.


3. Marsupella aquatica (Lindenberg) Schiffner, Sitzungsber. Deutsch. Naturwiss.-Med. Vereins Böhmen “Lotos” in Prag 44 (n.s. vol. 16): 267. 1896

Jungermannia emarginata var. aquatica Lindenberg, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 14(Suppl.): 75. 1829; Marsupella emarginata var. aquatica (Lindenberg) Dumortier


Plants erect, rigid, robust, to 50--120 x 1.5--3 mm (in terrestrial fo. pearsonii often just10 mm), in dull, dirty green to blackish red in exposed sites, reddish or carmine pigmented (fo. pearsonii), shoots usually simple except for subfloral innovations. Stem 250--320 x (300--)350-420 \um in diameter, without distinct hyalodermis, with cortical layer of slightly thick-walled, not strongly hyaline cells, intracortical cells in 2--4 layers smaller yellowish, considerably thick-walled, cells of medulla pellucid but slightly thick-walled. Rhizoids few, colorless to light violet. Leaves distant to approximate, towards apex contiguous, sheathing the stem, patent to suberect, rotundate, often slightly wider than long, averaging 1.1--1.3 as broad as long, usually with both margins reflexed, rarely plane, shallowly 2-lobed.  Sinus lunate or obtusely angulate, rarely more than 0.1 of the leaf length. Lobes wide, obtuse to rounded. Leaf cells at margins 10--12(--14) /um, in midleaf 0--28 x 18--22 /um  with distinct to bulging and confluent trigones, bistratose near base. Oil-bodies 2(--3) per cell, granular-botryoidal, 9--15 x 5--10(--12) /um. Sexual condition dioicous. Male plants with 3--5 pairs of imbricate bracts that are slightly lager than leaves, 2--5(--6) antheridia per bract. Female plants with subfloral innovations, bracts larger than leaves, Perigynium high, perianth hidden in bracts, its mouth subentire. [Spores large, to 20 /um wide, verruculose, not seen in N.America].

In beds and at the edges of swift, often cascading mountain brooks, also in shallow standing water in acidic lakes, often submerged, probably widespread in mountains of western and eastern North America, known from Greenland; B.C., Nfld., N.S.; Alaska, Maine, N.H., N.Y., Wash.; Eurasia; Atlantic Islands (Azores, Iceland, Faroes Islands).


Marsupella aquatica is probably under-recorded because it is often included in M. emarginata as var. aquatica. Marsupella emarginata is similar to M. aquatica in appearance but this last differs in (1) more rigid shoots that are usually much longer; (2) color of plants usually dirty green to blackish red; (2) more rigid and wider leaves that are usually broader than long and divided less than 0.2 leaf length; (3) rounded to obtuse lobes; (4) more prominent, often bulging and confluent trigones; (5) the less sharply differentiated hyalodermis of the stem.


4. Marsupella arctica (Berggren) Bryhn & Kaalaas, Bryoph. Itin. Pol. Norv. 11: 26. 1906


Sarcocyphos emarginatus var. arcticus Berggren, Kongl. Svenska Vetensk. Acad. Handl. (n.s.) 13(7): 96. 1875, as Sarcoscyphus; Marsupella groenlandica C.E.O.Jensen


Plants usually strongly pigmented, reddish brown to blackish, rarely dull green, ca. 10--40(--60) mm, erect to suberect. Stems 180--220 /um in diameter, in cross section with poorly developed hyalodermis of one row of slightly larger cells that are ca. 1.5--3 times larger than the intracortical, which are thick-walled with brownish walls; cells of medulla thin-walled, collenchymatous, approximately the same size as cells of hyalodermis; stolons often present in old parts of stem. Rhizoids colorless or reddish, rare. Leaves equal size throughout, almost hemispheric, spoon-shaped concave, almost rotundate, ca. 500 /um long and wide.  Sinus absent or to 0.1 the length of leave, margins incurved, often decolorated. Leaf cells at margins 10--15 /um, at midleaf 20--24(--27) x 18--21 /um, usually with bulging trigones. Oil-bodies 2(--3) per cell, large 5--9(--11) x 9--15 /um. Sexual condition dioicous. Androecial bracts in 2--4 to 5--7 pairs, similar to leaves. Both gynoecia and sporophytes unknown in North America.


Temporarily submerged at the edges of shallow pools, lakes, in seepages, along streams under permanent snow fields, peaty soil between rocks irrigated by permanent snow fields; near sea level up to 400 m elevation; Greenland; Nun., N.W.T., Que.; Alaska; Europe (Scandinavia, Scotland); N Asia, China.


5. Marsupella boeckii (Austin) Lindberg ex Kaalaas, Nyt Mag. Naturvidensk. 33: 409. 1893


Sarcocyphos boeckii Austin, Bull. Torrey Bot. Club 3(3): 9. 1872, as Sarcophus


Plants green, brownish green, to dark red and reddish brown, filiform, to 10 x 0.2--0.5 mm. Stems ca. 75--100 /um in diameter, with well-defined cells of hyalodermis. Rhizoids absent on main shoots or very few, colorless. Leaves distant to very distant, patent to suberect and erect, concave, with often incurved lobes, suborbicular to semicordate, usually wider than long, rarely as wide as long or longer than wide. Sinus acute to more or less rounded, reaching to 1/3--1/2 leaf length. Lobes broadly triangular, subacute or acute, when acute then ending in one or less often 2 mostly subisodiametric cells ca. 12--14 x 12--15 /um. Leaf cells at margins in 1--3 rows with incrassate walls, ca. (10--)11--12 /um, not decolorated, in midleaf slightly larger, ca. 13--18 /um, more or less thick-walled with distinct but usually small trigones; cuticle smooth to weakly papillose. Oil-bodies colorless, granular-botryoidal, 2(--4) per cell, ellipsoidal to ovoid, 5--8(--11) x 3--5 /um. Sexual condition dioicous. Male bracts in 2--5 pairs, imbricate, much larger than adjacent leaves, margins edentate; antheridia 1--3 per bract, antheridial stalk one seriate. Female inflorescence to 0.8 mm wide, bracts inflated, much larger than sterile leaves, with edentate margins. Capsule spheric, 2-stratose; elaters 6--7 /um, 2-spiral. Spores reddish brown, 15-18 /um. 


Arctic-montane, mostly oceanic and suboceanic, over thin layers of peat-covered or shaded acidic rocks, both in pure mats or mixed with Marsupella sphacelata, Gymnomitrion concinnatum, Lophozia sudetica; from near sea level to 900 m elevation; Greenland; Maine; Europe; Asia (Bhutan, Japan, Russia).


The only species of Marsupella that may be confused with M. boeckii is M. stableri. The latter differs from M. boeckii in appressed, more or less canaliculate leaves that are mostly longer than wide vs. patent to suberect and erect subqudrate or wider than long leaves in M. boeckii, and dentate or crenulate margins of male and female bracts. From Cephaloziella species with which it is often confused, M. boeckii differs in cells of leaves with usually distinct trigones, which do not occur in Cephaloziella, and different male and female inflorescence and stem anatomy.


6. Marsupella bolanderi (Austin) Underwood, Zoë 1(12): 365. 1891 E F


Sarcocyphos bolanderi Austin, Bull. Torrey Bot. Club 3(3): 9. 1872, as Sarcoscyphus


Plants small, erect, (2--)2--5 x (0.3--)0.4--0.6(--0.7) mm, green to reddish brown and brick colored especially distal part and margins of leaves, often with vertical shoots from horizontal stems, with numerous stolons. Stem in cross-section rounded, ca. 170--190 /um, with 2(--3) layers of large cells with more or less brown-colored thin walls with distinct but small trigones, ca. (15--)17--20(--25) /um, gradating to relatively small thin, yellow walled cells of medulla that are ca. (7--)8--12(--15) /um. Rhizoids numerous over ventral part of stem, long, colorless. Leaves spreading, wider than long, wide/length ratio ca. 1.1--1.35, quadrate-rotundate, 325--430 x 350 --500 /um, slightly larger below gynoecia, smaller proximally. Sinus obtuse, relatively deep, 0.3--0.4 leaf length. Leaf lobes narrowly ovate-triangular, obtuse to rounded, margins never reflexed. Leaf cells isodiametric and (18--)20--25 /um, or longer than wide and 30--34 x 22--25 /um in the middle and slightly smaller in lobes and along the margins, down to (15--)17--20 /um along margin, trigones small; base of leaves and especially bracts 2-stratose. Sexual condition dioicous. Female plants usually in separate patches; with relatively low apical Perigynium with perianth hidden in the bracts that much larger than caulinate leaves, 600--100- x 500--700 /um. Otherwise unknown.


On exposed rocks, damp sandstone outcrops, on sandstone hills; low to medium elevation; Calif., Oreg., Wash.


Marsupella bolanderi is endemic to the western United States, from Washington to California. It can be confused with small specimens of M. emarginata, it differs from the latter in (1) size smaller, not exceeding 5--6 mm whereas in M. emarginata even smaller forms are at least 10--15 mm; (2) leaves not pectinately inserted and oriented in M. bolanderi; (3) lobes narrowly triangular vs. broadly triangular in M. emarginata; (4) sinus never acute and relatively deep, reaching 03--0.4 leaf length vs. acute and usually not more than 0.2 -- 0.25 leaf length in M. emarginata; (5) margins of leaves never reflexed vs. narrowly reflexed in M. emarginata; and (6) absence of the thick-walled intracortical cell layers characteristic of M. emarginata. From small terrestrial forms of M. sphacelata it differs in (1) color red-brown or brick-colored vs. green to black-brown scorched, never red in the latter; (2) sinus of leaf never acute vs. acute and often closed sinus in M. sphacelata; (3) leaves spreading vs. leaves sheathing at base in M. sphacelata; and (4) absence of thick-walled intrarcortical cell layers characterestic of M. sphacelata. 


7. Marsupella condensata (Ångstrom ex C. Hartman) Lindberg ex Kaalaas, Nyt Mag. Naturvidensk. 33: 420. 1893


Gymnomitrium condensatum Ångstrom ex C.Hartman, Handb. Skand. Fl. (ed. 10) 2: 128. 1871


Plants more or less glossy, reddish brown, chestnut-brown to almost black, threadlike, terete, 5--30 x 0.17--0.3(--0.5) mm. Rhizoids frequent proximally, rare in distal part but sometimes on base of marsupium, colorless. Leaves erect to erect-spreading, moderately imbricate, but mostly not quite hiding the stem, distinctly concave, ovate-rotundate, 350--420 /um.Sinus  lunate or more rarely rotundate0.2--0.25 leaf length. Lobes triangular, acute and incurved at apex. Leaf cells at margins  (8--)10--18 /um, not or rarely decolorated just at the tip, in midleaf 16--22 x14--18 /um, trigones small to large but not bulging; cuticle smooth. Oil-bodies almost smooth, 2--3 (--4) per cell, ovoid to ellipsoidal, 7--11(--13) x 4--6 /um. Sexual condition dioicous. Male branches similar to sterile ones, bracts slightly larger than leaves, slightly ventricose at base, antheridia (2--)3--4 per bract. Female branches clavate, dorsiventrally compressed; bracts very broadly ovate-triangular, wider than long, 2-lobed for 0.15 their length, strongly concave. Perigynium short (0.25--)0.3--0.5 the height of the free perianth. Perianth with irregularly crenulate-denticulate mouth, cells at mouth fingerlike, 24--38 x 15--17 /um. Capsule spheric, 2-stratose; epidermal cells with coarse nodular thickenings; elaters 7--8 /um, 2-spiral. Spores almost smooth, reddish or yellow-brown, 10--13 /um. 


Arctic-montane mostly oceanic and suboceanic, restricted to non-calcareous sites continuously irrigated by water from melting snow; Greenland; B.C., Yukon; Alaska; Europe; Asia; Atlantic Islands (Iceland, Svalbard).


Marsupella condensa is easily distinguished from other Marsupella species, apart from M. apiculata. From the last it differs in (1) leaves not cuspidate, ending in one celled apiculus; ( 2) margin of leaves not decolorate; (3) sinus lunate or rotundate vs. acute in Marsupella apiculata; and (4) shoots with less strongly imbricate leaves.


8. Marsupella emarginata (Ehrhart) Dumortier, Recueil Observ. Jungerm., 24. 1835


Jungermannia emarginata Ehrhart, Hannover Mag. 22: 141. 1784


Plants erect, (10--)20--50 x 0.5--2.5 mm, green to reddish brown, red or carmine, sparingly branched, often with stolons, frequently with subgynoecial innovations. Stem in cross-section with one-layered hyalodermis and 3--4 layers of tangentially-flattened thick and yellow walls, smaller intracortical cells medulla of thin-walled cells. Rhizoids few, mostly restricted to stolons and bases of shoots, colorless, rarely vinaceous. Leaves approximate to contiguous, at base more or less sheathing the stem, spreading, often pectinately inserted, more or less equal in size, slightly larger below gynoecia, rotundate to quadrate-rotundate. Sinus  angular to acute, 0.2--0.3 of the leaf length. Lobes broadly triangular, obtuse or obtusely pointed; margins narrowly reflexed. Cells at margins 11--13/um with more or less thickened walls  and (16--)18--23 /um in the middle with large trigones; . Oil-bodies finely granular, 2(--3) per cell, ovoid to ellipsoidal, 6--8 x 4--6 /um.  Sexual condition dioicous; male and female plants usually in separate patches. Male plants with 3--5(--10) basal pairs of bracts that are shallowly 2-lobed, saccate at base, 2--5(--6) antheridia per bract, androecia apical, becoming intercalary. Female plants with high, fleshy apical Perigynium and perianth hidden in bracts, bracts similar to but slightly larger than leaves, perianth mouth subentire, irregularly lobed or divided into narrow lobes. Capsule spheric; walls (2--)3-stratose, elaters with 2(--3)-spiral. Spores 9--13(--20) /um, red-brown, distinctly verruculose.


Mountain pioneer on wet and damp acidic rocks, sometimes on decaying wood near swiftly running water, particularly at the edges of mountain streams, waterfalls and cascades where humidity is high; low to medium elevations; probably throughout mountains of western and eastern North America, Greenland; Alta., B.C.,  N.B., Nfld. and Labrador, N.W.T., N.S., Nunavut, Ont.,  Que., Yukon;  Alaska, Calif., Colo., Conn., Ga., Ky., Maine, Mass., Mich., Minn., Mont., N.H., N.Y., N.C., Oreg., Pa., Tenn., Vt., Va., Wash., W.Va.; Europe; Asia (China, Japan, Russia); tropical Africa; Atlantic Islands (Azores, Canary Islands, Faroes, Iceland, Madeira);


Marsupella emarginata is one of the most widespread species of the genus. The species can be confused with M. aquatica, M. bolanderi, M. paroica and M. sphacelata. For the differences with M. aquatica, M. bolanderi and M. paroica see those species. It differs from the M. sphacelata in (1) color green, olive-green to reddish brown, whereas M. sphacelata is usually bright or dull green to black-brown and scorched; (2) stiffer, pectinate leaves vs. soft and lax in M. sphacelata; (3) leaves with reflexed or recurved dorsal and most ventral margins, also in bracts, whereas the margins are plane in M. sphacelata; (4) less deeply and sharp 2-lobed leaves, sinuses in M. emarginata rarely reach 0.3 leaf length and they reach 0.5(--0.6) leaf length in M. sphacelata; and (5) mostly subacute to obtuse lobes vs. these mostly rounded  in M. sphacelata). 



9. Marsupella funckii (F. Weber & D. Mohr) Dumortier, Recueil Observ. Jungerm, 24. 1835


Jungermannia funckii F. Weber et D. Mohr , Bot. Taschenb. (Weber) 1807: 422. 1807


Plants deep green-brown, fuscous to blackish, erect or suberect, shoots ca (3--)5--10(--15) x 0.35--0.70(--0.80) mm in sterile shoots, much wider in gynoecial area to 1.35 mm broad; stolons few or absent. Stems (9--)10--12(--13) cells high, hyalodermis absent, cortical cells thick-walled, one layer larger than adjacent, intracortical cells in 2--3 layers similar to cortical but somewhat smaller grading to smaller and thin-walled cells of medulla. Rhizoids few, near the base. Leaves pectinately spreading, on sterile shoots nearly subequal size, rounded-quadrate to broadly ovate, bases rounded or cordate, with plane margins. Sinus acute, not reflexed, to 0.35--0.45 of leaf length.  Lobes triangular, acute. . Cells medium-sized, marginal ca. (11--)12--15 /um, in midleaf  16--20 x 14--18 /um with distinct trigones, cuticle smooth. Oil-bodies finely granular, 2(--3) per cell, ovoid to ellipsoidal, 6--8(--10) x 3--5 /um. Sexual condition dioicous. Male bracts in 2-several pairs, larger than adjacent leaves; antheridia 2 per bract. Female bracts much larger than leaves, Perigynium low, about equal in height to free perianth, which is well developed but much shorter than bracts. Capsule spheric, with 2-stratose wall of epidermis, inner cells of capsule with nodular thickenings. Spores pale brown, 7--9 \um, nearly smooth.


Montane species, on moist soil-covered rocks in mountains; 1700--1800 m; Tenn.; Europe; Asia (Turkey, Japan, South Siberia), Atlantic Islands (Azores, Iceland, Macaronesia).


Marsupella funckii can be confused with terrestrial forms of M. sphacelata from which it differs in (1) stem anatomy, having 2--3 layers of intracortical cells slightly different from the cortical cells, whereas in M. sphacelata the hyalodermis is subtended by 3--4 layers of much smaller and deeply brownish pigmented cells; (2) acute leaf lobes, which in M. sphacelata are broadly rounded or obtuse; (3) smaller leaf cells; (4) smaller size of terrestrial plants; and (5) different color of shoots. Small plants of M. emarginata differ in size (much bigger), less deep divided leaves, color being mostly with different tints of red; recurved margin of leaves, and stem anatomy.


10. Marsupella paroica R. M. Schuster, Bryologist 60: 145. 1957


Plants green to somewhat brownish tinged, never purplish or black, erect, sterile shoots (5--)10--15 x 0.8--1.2(--1.6) mm. Stems 13--16 cells high, with one row of large thin-walled hyaline outer cells and 2--3 rows of much smaller inner cortical cells with very thick brownish or yellowish brown walls; cells of medulla thin-walled and as large or slightly larger than outer cells. Rhizoids few, colorless. Leaves slightly succubously inserted, broadly ovate, widest at or somewhat below the middle, slightly longer than wide, distinctly unequally 2-lobed. Sinus subrectangular to subacute, rarely acute 0.15--0.2(--0.25) of the leaf length. Leaf lobes subobtuse to acute or subacute with margins from the distal third of leaves often (but not always and usually only on one side) narrowly reflexed. Cells of margins of leaf in one row much smaller, ca. 10--12 /um, quadrate to shortly-rectangular, not decolorated, inner cells equally thick-walled ca. 20 --25 x 18--22 /um, with bulging trigones. Oil-bodies finely granular, 2(--3) per cell, spherical to ovoid and ellipsoidal, 9--12 x 5--7 /um. Sexual condition paroicous. Male bracts below gynoecia in 2--4(--5) pairs imbricate with distinctly saccate base, wider than long and larger than leaves, with 1--3 long, 2-seriate stalked antheridia per bract. Perianth terminal, hidden in suberect shallowly 2-lobed bracts. Sporophytes with subspheric capsule, inner cell walls with generally 4--7 nodular thickenings, epidermal layer with 1--2 nodular thickenings per longitudinal wall and 0--1 per transverse wall. Spores brown, 10--13 /um, elaters 2-spiral.


Humid areas, almost always in deep shade on rather dry to damp and moist rocks, sometimes on decaying wood near flowing water especially along cascades, waterfalls, swift mountain streams, usually on the heights; 1600--1900 m; Ontario; Georgia, Minn., N.C., S.C., Tenn., Va.; Mexico.


Marsupella paroica is restricted to the Southern Appalachians and Mexico. It can be confused with M. emarginata from that it differs in (1) paroicous inflorescence; (2) generally narrower lobes; (3) in general smaller shoots; (4) absence of the purplish red pigmentation typically found in sun forms of M. emarginata; (5) different capsule wall anatomy. From Marsupella sparsifolia, which is somewhat similar in size and also paroicous, it differs in (1) leaves more spreading, shallow and less acute, unequally divided; (2) different color of shoots that never are brown to brownish black; and (3) leaf margins often reflexed.  


11. Marsupella sparsifolia (Lindberg) Dumortier, Bull. Soc. Roy. Bot. Belgique 13: 128. 1874


Sarcocyphos sparsifolius Lindberg, Not. Sallsk. Fauna Fl. Fenn. Forh. 9: 280. 1868, as Sarcoscyphus


Plants golden brown to chestnut and blackish brown, rather scorched, more or less shiny, erect or suberect, often in pure mats, shoots 6 --10(--12) x 0.45--0.60 mm, to 1.5 mm wide in sterile shoots, in gametangial area reaching 8 mm broad; stolons few or absent. Stems (9--)10--12(--13) cells high, ca. 125--150 /um in diameter with poorly defined hyalodermis, intracortical cells in 2--3 layers similar to the cortical, thin-walled, ca. 17--22 /um in older parts, these contrasting with ill-defined small thin-walled cells of medulla that are (10--)12--15 /um. Rhizoids few, colorless or rose to reddish brown. Leaves spreading, erectopatent to squarrose, subequal in size but slightly larger when fertile, concave or slightly sheathing the stem (250--)400--800(--900) x (250--)400--700 /um, subquadrate or slightly longer than wide.  Sinus open, from acute to rectangular or narrowly rounded, 0.25--0.3 of leaf length.  Lobes triangular or broadly triangular acute or subacute to blunt with margins plane and not decolorated.. Cells of the midleaf 17--25 x 14--17 /um with distinct trigones, cuticle smooth. Oil-bodies granular papillose, 2(--3) per cell, subspheric to ovoid and ellipsoidal, 5--7(--11) x 4--6 /um. Sexual condition paroicous. Male bracts in 2--3 pairs below gynoecia, slightly inflated at base and slightly larger than adjacent leaves with 1--3 antheridia per bract, with 2-seriate stalk. Female bracts larger than leaves, wider than long, to 1350 x 1400 /um, Perigynium low. Sporophytes with spheric capsule, 2-stratose walls of epidermis and inner cells of capsule with nodular thickenings. Spores brown, 10--15 \um, nearly smooth; elaters mostly 2(--4)-spiral, 10--12(--14) /um wide with band 2--3 /um.


Arctic-montane, on soil over rocks, on moist steep stabilized boulder slopes, on wet rocks along and in streams, rock outcrops in forests, on exposed face, moist seepages, in cliff crevices, damp outcrop faces, on banks of glacial streams; 900--1500 m; Greenland; Alta., BC; N.S., Ont., Que.; Calif., Mich., Mont., N.H., Wash.; c,s Africa; Atlantic Islands (Azores, Faroes); Pacific Islands; Antarctic (subantarctic islands).


Marsupella sparsifolia is almost always fertile and can be confused with other Arctic-montane paroicous species, particularly with M. sprucei from which it differs in (1) being larger; (2) having more regularly pectinate shoots; (3) more strongly spreading, even squarrose leaves that are not very much smaller and never scale-like nor semi-appressed proximally; (4) larger leaf cells; (5) suberect concave sheathing base and different color of shoots, and (6) often purplish red rhizoids. For differences among them and another paroicous species in the flora, M. paroica as well as from M. sphacelata see these species.


12. Marsupella sphacelata (Gieseke ex Lindenberg) Dumortier. Recueil Observ. Jungerm., 24. 1835


Jungermannia sphacelata Giesecke ex Lindenberg, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 14(Suppl.), 76, pl. 1. 1829; Marsupella sullivantii Evans; M. jorgensenii Schiffner


Plants erect, (8--)10--30 mm in terrestrial forms and up to 40--70 mm high in aquatic forms, 0.5--3 mm wide, bright or dull green, brown distally and on leaf lobes, to black-brown, appearing scorched, sparingly branched, with numerous stolons in aquatic forms. Stem relatively soft, in cross-section with one- layered hyalodermis collapsed with age and (2--)3--4 layers of deeply brownish pigmented, moderately thick-walled intracortical cells and rather small leptodermous medullar cells. Rhizoids few, colorless or often vinaceous. Leaves gradually larger proximally, with bases sheathing the stem, subquadrate or slightly wider than long (in aquatic forms) or longer than wide (in terrestrial forms), margins plane. Sinus acute, open to and closed proximally, 0.3--0.5(--0.6) leaf length. Lobes usually broadly rounded (rarely subacute to obtuse) from suberect to incurved, squarrose and spreading. Cells in terrestrial forms18--25 x (15--)18--22 /um, in aquatic forms larger to 23--28 x 20--22 in the middle of the leaf, and (11--)15--18 on margins, trigones distinct, but in aquatic forms rather small or absent. Oil-bodies granular papillose, 2--3 per cell, spherical to ovoid and ellipsoidal, 6--9(--12) x 3--5 (--7) /um. Cuticle verruculose-striolate. Sexual condition dioicous. Male plants with imbricate bracts, oblong-quadrate, divided 0.25--0.35 of the leaf length, with 2--3 long-stalked antheridia per bract. Female plants with apical gynoecium and high fleshy Perigynium; Female bracts similar but slightly larger than leaves, perianth shorter than bracts. Capsule spheric, yellow-brown; walls 3-stratose. Spores 9--12 /um, yellow-brown, nearly smooth.


Montane pioneer on damp acidic rocks, on banks of stony streams, rarely in standing water; medium to high elevation (2700 m alt. in California); Greenland; Miquelon; Alta., B.C., Nfld., Que., Yukon; Alaska, Calif., Colo., Conn., Ga., Idaho, Ky., Maine, Mass., Miss., Mont., N.H., N.Y., N.C., Oreg., Tenn., Vt., Va., Wash., W.Va.; Europe; Asia (Caucasus, Far East of Russia, Japan, s Siberia); Atlantic Islands (Azores, Iceland).


Marsupella sphacelata has a holarctic, mostly boreo-montane, circumpolar distribution. It is widespread in the mountains of subarctic regions. The species can be confused with M. funckii, M. sparsifolia, M. emarginata. It differs from the latter in (1) color green, dull green to black-brown, usually brownish distally and characteristically appearing scorched, whereas M. emarginata is often reddish brown to carmine-red (red pigmentation almost always present), never scorched; (2) leaves sheathing basally but patent distally vs. almost vertically spreading and slightly or not sheathing in M. emarginata; (3) more deeply lobed leaves, often up to 0.3--0.4 of leaf length, whereas in M. emarginata the sinus does not exceed 0.25 of leaf length; and (4) plane, never reflexed margins vs. always reflexed at least in some leaves in M. emarginata. Terrestrial forms can be confused with M. sparsifolia because of small size and similar black-brown color. They differ from M. sparsifolia in (1) dioicous inflorescence; (2) rounded or blunt leaf lobes vs. acute or subaute leaf lobes in M. sparsifolia; (3) stem anatomy, having a clearly distinguishable hyalodermis vs. ill-defined hyalodermis in M. sparsifolia.


Marsupella sphacelata is a very variable species. Apart from lax and large aquatic forms, several small terrestrial forms can be distinguished, some of which were described as separate species. Marsupella sullivantii Evans was treated as M. sphacelata fo. media by R. M. Schuster (1974). It has leaves bifid 0.3--0.45 their length widest above or near the middle and reflexed and rounded to blunt lobes. It is the most widespread form. Another small terrestrial form was described as Marsupella jorgensenii Schiffner, and treated as M. sphacelata fo. joergensenii (Schiffner) R. M. Schuster (R. M. Schuster 1974). This form has more or less squarrose ovate to broadly ovate widest near base, more deeply divided to 0.35--0.45 of their length leaves, and leaf lobes triangular to acute or subacute, never broadly rounded.


13. Marsupella spiniloba R. M. Schuster & Damsholt, Phytologia 63 (5): 326. 1987


Plants brownish black, small, shoots flattened distally, 2.5--5 x 0.2--0.4 mm, branches lateral-intercalary. Stems 0.1--0.2 mm in diameter, in cross section 8--10 cells high, with poorly differentiated cortical cells, cells on dorsal side of stem relatively large, ca. 14--18(--20) x 18--22(--25) /um; stolons and geotropic flageliform axes numerous. Rhizoids numerous, colorless. Leaves in sterile shoots pectinate-distichous, contiguous to imbricate, patent to squarrose, towards shoot apex canaliculate, broadly ovate, without decolorate plane margins, V-shaped narrowly acute at base 0.25--0.35 leaf length, sharply pointed, acuminate, with one or both divergent tapered apices ending in (1--)2--3 superposed cells. Cells of leaf middle as long as wide, 10--13(--15) /um, marginal cells clearly different from innermost cells, (7--)9--11(--12) /um, trigones relatively large; cuticle smooth. Oil-bodies large, mostly 2 per cell, spheric to ellipsoidal, mostly 2--3 /um, but also 4 x 7 μm, finely granular. Sexual condition paroicous. Female bracts larger than the leaves similar in shape, perianth shorter than the bracts; perianth mouth crenulate. Male bracts in 1--2 pairs broadly ovate, slightly larger than leaves, antheridial stalk 2-seriate. Seta ca. 100--105 /um. Elaters 3-4-spiraled. Spores granulate-vermiculate, light brown 15--16.5 /um.


Arctic-alpine, occurring on open soil or rocky slopes; low to moderate elevations; S. Greenland; Alaska; Europe (Scandinavia).


Marsupella spiniloba is a poorly known, problematic taxon. It can be confused with M. sprucei, from which it differs in (1) acuminate, divergent leaf lobes, vs. acute to subacute in M. sprucei; (2) distinct habitus of vegetative shoots that have flattened apices in M. spiniloba and not in M. sprucei; and (3) smaller leaf cells of leaf margins contrasting with larger medial cells in M. spiniloba, whereas they are not clearly distinct in M. sprucei.


14. Marsupella sprucei (Limpricht) Bernet, Cat. hép. Suisse, 33. 1888


Sarcocyphos sprucei Limpricht, Flora 64 (5): 72. 1881, as Sarcoscyphus; Marsupella ustulata var. sprucei (Limpricht) R. M. Schuster


Plants dark green to purplish and brownish black, erect from creeping rhizomatous stem, small-leaved at the base and becoming more or less rapidly larger-leaved distally, sterile shoots 3--5 x 0.30--0.45 mm, in gametangial area reaching 0.80 mm broad, often fasciculate with numerous lateral-intercalary branches often flagelliform and then more or less small-leaved, and with small-leaved stolons and geotropic microphyllous to leafless rhizioidous axes. Stems ca. 100 /um in diameter, often flagelliform with poorly defined hyalodermis. Rhizoids rare on clavate fertile branches but numerous on creeping or descending flagelliform shoots, colorless. Leaves on sterile branches more or less spreading, imbricate and small near shoot basis, on fertile shoots erect to erect-appressed and significantly larger in gametangia area, concave, broadly ovate to ovate-quadrate, widest near middle or near base, on sterile shoots 240--275 x 250--278 /um. Sinus acute, 0.2--0.3(--0.35) leaf length.  Lobes acute or subacute, often ending in 2 superposed cells, margins of leaves plane, not decolorated, sinus acute. Cells vary greatly in different forms, the marginal variously (8--)9--13(--16) to (13--)15--20 x (12--)14--25 in mid leaf. Oil-bodies granular papillose, 2 (--3) per cell, subspherical to ovoid and ellipsoidal, finely granulate, 5--9 x 4--6 /um. Sexual condition paroicous. Male bracts in 1--2(--3) pairs just below gynoecia, concave, imbricate, wider near base, much larger than adjacent leaves with subacute or blunt lobes, 1--3-androus. Female bracts erect, not or cordate at base, much larger than leaves, broadly ovate with usually blunt lobes. Perigynium to 0.3--0.4 height of well-developed perianth and hidden in the longer bracts, mouth of perianth crenulate. Sporophytes with spheric, reddish brown capsule, epidermal layer with few very coarse, stalked nodular thickenings mostly on longitudinal walls and much weaker nodular thickenings on inner layer. Elaters 2 spiral, 6--8 /um wide. Spores smooth light reddish brown, 7--9 \um..


Arctic-montane, on exposed bare soil on soil over rocks, on bare and on moist peaty soil in tundra, in late lying snowbeds, cliff crevices, bank of glacial stream; 850--1750 m; Greenland; BC; N.S., Nunavut, Que.; Alaska, Calif., Maine, N.H., Oreg., Vt., Wash., Wyo.; Europe; Asia (Siberia, China); Atlantic Islands (Faeroes, Iceland, Madeira); Pacific Islands (New Zealand), Antarctic (subantarctic islands).


Marsupella sprucei is almost always fertile and can be confused with another Arctic-montane paroicous species, Marsupella sparsifolia as well as with poorly known M. spiniloba. For differences see these species. Marsupella sprucei is a very variable species. Apart from the type variety Marsupella sprucei var. ustulata (Limpr.) Damsh. has been distinguished in the flora. It differs in smaller leaf cells and spores, cordate at base male and female bracts without acute lobes.


15. Marsupella stableri Spruce, Rev. Bryol. 8: 96. 1881

Marsupella boeckii var. stableri (Spruce) R. M. Schuster


Plants from pinkish green, rosy to dark red, purplish red and almost black, rarely bright green or brownish green and reddish brown, filiform, 5 x 0.12--0.2 mm, grooved along visible antical surface, stems with well-defined hyalodermis, cross-section of stem 12 cells high, cells on dorsal side of stem 20 x10--12(--15) /um, cells of hyalodermis in 2 rows large, thin-walled, 15--17 x 12--15 /um, gradually smaller, cells of medulla thick-walled, (5--)7--10 /um. Rhizoids absent on main shoots or very few, colorless. Leaves from slightly distant to imbricate, suberect or erect-appressed, canaliculate, scale-like proximally, distinctly longer than wide. Sinus acute 0.35--0.5 leaf length. Lobes narrowly triangular, acute ending in one often elongated cell, 15--17 x 8 /um or 2 almost isodiametric superimposed cells, and then ca. 7--9 /um. Leaf cells along margins in 1--3 rows with thick walls, ca. 7--8 x 10--13 /um, not decolorate, medial cells slightly larger, ca. 12--18 x 10--13 /um, thin-walled, with distinct but usually small trigones; cuticle smooth to rather distinctly papillose. Oil-bodies 2(--3) per cell, spherical to ovoid, 3-8 x 3--6 /um. Sexual condition dioicous. Male inflorescence to 0.5 mm wide, capitate, bracts in 1--5 pairs, imbricate, much larger than adjacent leaves, lobes spreading, margins crenulate; antheridia one per bract, antheridial stalk one seriate. Female inflorescence to 0.8 mm wide, bracts inflated, larger than sterile, bracts with dentate margins, perianth hidden in bracts, cells of mouth almost isodiametric, ca. 25 x 20 /um. Perigynia well developed. Sporophytes unknown in flora area [walls of epidermal and inner cells of capsule valves with narrowly based nodular trigones. Spores 10--15 /um].


Oceanic species of restricted range, rocks and damp open outcrops in bogs and on open boggy slopes, stones in opening of bogs; low to moderate elevations; Alaska; B.C.; Europe (Great Britain).


Marsupella stableri is found in pure mats or mixed with M. sphacelata or M. emarginata. The only species of Marsupella that can be confused with M. stableri is M. boeckii. Marsupella stableri differs from M. boeckii in its appressed, more or less canaliculate, imbricate leaves, which are longer than wide, and the crenulate margins of male bract and dentate margins of female bracts. The treatment of J. Váňa et al. (2010: 47) is followed here, which recognized M. stableri as a distinct species while R. M. Stotler and B. J. Crandall-Stotler (2017) followed R. M. Schuster (1974: 102) in placing Marsupella stableri in synonymy of M. boeckii.



3. NARDIA Gray, Nat. Arr. Brit. Pl. 1: 694. 1821, as “Nardius * [For S. Nardi, an Italian abbot.]


Marie L. Hicks



Plants prostrate, ascending when crowded, forming mats, green to reddish or brownish.  Stems thick and fleshy; branching intercalary or terminal; cortical cells thin‑walled, 32‑‑40 ´ 16‑‑28 /um, sometimes reddish tinged, not distinctly differentiated from slightly longer medullary cells, 35‑‑60 x 16‑‑40 /um; rhizoids scattered along ventral stem in irregular fascicles, some from leaf bases.  Leaves succubous‑oblique, broad, as wide as long or wider, entire, with or without 2-lobed apex; leaf cells rounded‑hexagonal with small to large trigones; oil bodies few, large, smooth or granular, opaque or hyaline.  Underleaves small, lanceolate to spatulate, often attached on one side to lateral leaf.  Specialized asexual propagation absent.  Sexual condition dioicous or monoicous.  Androecia terminal, becoming intercalary; bracts similar to leaves, larger, not or slightly modified; antheridia 1‑‑3 per axil, stalks 2‑seriate.  Gynoecia terminal on main shoots; bracts, inserted on fleshy perigynium, unmodified or shallowly lobed, large in comparison to perianth and concealing it; bracteole present, subulate to lanceolate; perianth short, conical, contracted to crenulate mouth; thickened stem forms a fleshy stem perigynium at base of perianth; calyptra developed atop perigynium; old  archegonia situated on calyptra.  Sporophyte foot imbedded in base of perigynium; seta 7‑‑8 cells in diameter; capsule globose to ovoid, 4‑valved, the walls 2‑cells thick; cells of exterior layer large, with nodular thickenings, inner layer smaller with semiannular bands;  elaters 150‑‑200  x 8‑‑10 /um, 2‑‑4 spiral. Spores 9‑‑24 /um.


Species 14 (8 in the flora), moist soil or humus: North America, South America (Brazil), Europe, Asia, Africa.


The genus Nardia is distinguished by the wide, entire to 2-lobed leaves, the lanceolate to spatulate underleaves that are sometimes narrowly connate with lateral leaves on one side.  The plants also have unspecialized androecia with bracts scarcely concealing antheridia.  Underleaves vary in size and may be vestigial on weak shoots or on proximal portions of stems.  They are best developed and should be searched for on apical parts of robust shoots.


SELECTED REFERENCES Schuster, R. M. 1969.  The Hepaticae and Anthocerotae of North America, Vol. 2. New York.  Hong, W.S.& J. Vana 2000. The distribution of Nardia in western North America, Lindbergia 25: 9--14.  Bakalin V.A. 2012. Nardia hiroshii Amak.---a new species for North American liverwort flora and the key to Nardia species in North Pacific. Arctoa 21: 87--90.


1.  Plant shoots 0.5 mm or less wide; leaves 2-lobed; trigones not distinct in leaf cells ..6. Nardia breidleri

1.  Plant shoots usually more than 0.5 mm wide; leaves entire or 2-lobed; trigones distinct in leaf cells.

2.  Plants paroicous, androecia beneath perianth, underleaves in sterile shoots vestigial to small, easily deciduous.

3.  Most leaves not lobed, a few leaves shallowly 2-lobed ... 4. Nardia geoscyphus

3.  All leaves 2-lobed ..............5. Nardia insecta

2.  Plants dioicous, androecia and gynoecia on separate plants, underleaves distinct to small, but then hyalodermis developed and leaves are laterally compressed to the stem.

4. Leaves 2-lobed.

5. Leaves shallowly 2-lobed, lobes obtuse; oil bodies 2--3 per mid leaf cell; underleaves spathulate … 3. Nardia lescurii

5. Leaves 2-lobed for 1/4--1/5 of leaf length, lobes usually acute; oil bodies 2--5 per midleaf cell;

6. Underleaves laciniate to narrowly triangular, perigynium not pendent … 8. Nardia hiroshii

6. Underleaves spathulate to triangular-lanceolate, perigynium short-pendent … 9. Nardia japonica

4. Leaves unlobed.

7..Oil bodies 1 per cell, coarsely granulate, present in ca. 30% of leaf cells; underleaves commonly connate with leaves in one side … 7. Nardia assamica

7. Oil bodies 1--4 per cell, homogenous, smooth to botryoidal, present in all (or nearly so) cells; underleaves (if present) triangular to subulata, not connate with the leaves .

8.  Leaves orbicular, marginal leaf cells only slightly smaller than median cells; stem hyalodermis absent ......... 1. Nardia scalaris

8.  Leaves reniform, marginal leaf cells distinctly smaller than median cells; stem hyalodermis present ...................2. Nardia compressa


1.  Nardia scalaris (Schrader) Gray, Nat. Arr. Brit. Pl. 1: 694. 1821


  Jungermannia scalaris Schrader, Syst. Samml. Krypto. Gewachse 2: 4. 1797


Plants with shoots 10‑‑30 x 1.5‑‑2.4 mm, prostrate, forming mats, light green to reddish brown.  Stems creeping or ascending when crowded, 360‑‑300 mm in diameter, branches few, terminal or innovating below perianths; rhizoids abundant, scattered along ventral stem, often with fascicles from leaf and underleaf bases, colorless or slightly brownish.  Leaves  contiguous to imbricate, slightly concave, erect to spreading, cuticle smooth, walls thin, trigones developed, sometimes bulging. Underleaves spreading, distinct, subulate to lanceolate, some narrowly connate on one side with lateral leaves, apices acute to acuminate.  Sexual condition dioicous.  Androecia intercalary, bracts in 3‑‑5 pairs, concave, imbricate, similar to leaves.  Gynoecia terminal on fleshy apex of main stem; bracts similar to and larger than leaves, 0.9‑‑1  x 1.2‑‑1.3 mm, ovate to reniform, concave, sometimes undulate or emarginate, connivent over perianth; bracteole subulate to lanceolate, larger than underleaves, narrowly connate to bracts; perianth short, ca. 300 /um, conical, hidden within the bracts, contracted to a crenulate mouth; perigynium fleshy, longer than perianth, 400‑‑600 mm, the base often tinged with red, bearing rhizoids.  Sporophyte capsule subglobose, dark brown; elaters brown, 2‑spiral. Spores 16‑‑18 /um, finely papillate, yellowish brown.  


Subspecies 2 (2 in the flora).


1.  Oil bodies glistening, hyaline, homogeneous, segmented with age into 2‑‑3 segments ..1a. Nardia scalaris subsp. scalaris


1.  Oil bodies opaque, botryoidal, made up of several droplets . 1b..Nardia scalaris subsp. botryoidea


1a. Nardia scalaris subsp. scalaris (Schrader) S. F. Gray


 Plants with shoots 10‑‑30   x 1.5‑‑2.4 mm.  Leaves circular to reniform in outline, about as wide as long or wider, 0.6‑‑0.9   x 0.7‑‑1 mm, entire with rounded apices, distal leaves occasionally retuse; median leaf cells 30‑‑35  x 24‑‑30 /um, marginal cells smaller, 20‑‑30 /um, oil bodies 2‑‑3 per cell, ovoid to ellipsoid, 8‑‑15  x 6‑‑7 /um, homogeneous, hyaline, glistening, becoming 2‑‑3 segmented with age.


Moist to dripping non‑calcareous rocks and along stream banks in Spruce‑Fir or Arctic; e, w, and s Greenland; B.C., Labrador, N.B. Nfld., N.S., Yukon; Alaska, Maine, N.C., Oreg., Tenn., Wash., Wyo.; Europe; Asia; Atlantic Islands (Iceland).


1b.  Nardia scalaris subsp. botryoidea R. M. Schuster, Hepatic Fl. N. Amer. 2: 862. 1969


Plants with shoots 10‑‑30 x 2‑‑2.5 mm.  Leaves circular to reniform in outline, up to 1.2  x 1.4 mm with some leaves emarginate, the indentation sharp, the lobes broadly rounded; median leaf cells 30‑‑40  x 28‑‑32 /um, marginal cells 28‑‑38 /um, oil bodies 2‑‑3(‑‑6) per cell, ovoid to ellipsoid, 10‑‑16  x 6‑‑9 /um, granular botryoidal, made up of numerous small droplets, opaque.


Soil over rock with seepage or on peat in bogs; w Greenland; N.S.; Tenn. (Great Smoky Mountains).


Fertile plants of Nardia scalaris subsp. botryoidea often have reddish undersides, especially near the bulbous perigynium and around the base of rhizoids.  Both subspecies are found in Tennessee above 1520 m in Spruce‑Fir forests.


2.  Nardia compressa (Hooker) Gray, Nat. Arr. Brit. Pl. 1: 694. 1821


   Jungermannia compressa Hooker, Brit. Jungermanniaceae pl. 58, 1816


Plants with laterally compressed shoots 20‑‑120 x 2‑‑3 mm, erect or sub‑erect, forming thick mats or turfs, green to reddish‑brown or purplish, often appearing scorched.  Stems fleshy, 250‑‑360 /um in diameter, branches few, intercalary from distal stems; rhizoids few, colorless, often absent near stem apex; hyalodermis present.  Leaves imbricate, erect‑appressed, orbicular to reniform, ca. 1‑‑1.8 x 1.2‑‑2.8 mm, with rounded, entire apices, slightly concave, shortly decurrent dorsally; median leaf cells 30‑‑40  x 25‑‑35 /um, marginal cells smaller, subquadrate, 18‑‑25 /um; cuticle smooth; trigones distinct, small to large and bulging; oil bodies  1‑‑3 per cell, ovoid to ellipsoid, 10‑‑14  x 7‑‑10 /um, shining, smooth, homogeneous or segmented with few segments.  Underleaves spreading, small, up to 0.5 mm, subulate to lanceolate, most developed at shoot apices, often vestigial on lower part of stem. Sexual condition dioicous.  Androecia terminal, becoming intercalary; bracts in 3‑‑4 pairs, similar to leaves.  Gynoecia terminal on main stems, bracts inserted on perigynium, larger and broader than leaves, reniform, exceeding length of perianth, hiding it; bracteole lanceolate, occasionally lobed, not connate to bracts; perianth conical, purplish, mouth crenulate; perigynium swollen, often purplish, continuous with stem, ca. 2 times longer than perianth.  Sporophyte capsule brown; elaters 2‑spiral. Spores 10‑‑15 /um, slightly papillate, reddish‑brown.


Wet rocks along streams or in peaty bogs, arctic-alpine; s Greenland; B.C.; Alaska, Wash.; Europe; Asia; Atlantic Islands (Iceland).


3.  Nardia lescurii (Austin) Underwood, Bull. Ill. State Lab. Nat. Hist. 2: 115. 1884


    Alicularia lescurii Austin, Hep. Bor.‑Amer. 4. 1873


Plants with shoots 15‑‑30 x 0.8‑‑1.8 mm, prostrate with ascending apices, in mats or thick patches, green to reddish tinged.  Stems soft and fleshy, 250‑‑350 /um in diameter; branches few, intercalary or terminal; rhizoids numerous, from base of leaves, underleaves and scattered along stems, colorless to slightly tinged with red.  Leaves approximate to imbricate, spreading, slightly concave, wider than long, 0.3‑‑0.7  x 0.4‑‑0.9 mm, shallowly 2-lobed with broad, obtuse lobes, the sinus less than 1/4 leaf length with ventral lobe slightly larger; median leaf cells 25‑‑40  x 28‑‑40 /um, marginal cells smaller (20‑‑24 /um); cuticle smooth to slightly verruculose, walls thin, trigones large, bulging; oil bodies 3‑‑5 per cell, ovoid to ellipsoid, 7‑‑10  x 10‑‑16 /um, granular.  Underleaves lanceolate with acute to acuminate apices, free or narrowly connate to lateral leaf on one side.  Sexual condition dioicous.  Androecia terminal, becoming intercalary; bracts in 6‑‑15 pairs, not differentiated from stem leaves, 2-lobed, not or only slightly concave with antheridia exposed in axils.  Gynoecia terminal on main stem, often with subfloral innovations; bracts inserted on perigynium, similar to but larger than leaves, 2--3-lobed, undulate, surpassing length of perianth, hiding it; bracteole ovate to lanceolate, free, apex acute to acuminate, occasionally with lateral tooth; perianth conical, short, 700‑‑1000 /um, mouth entire or irregularly lobed and crenulate; perigynium fleshy, as long as perianth or longer, 700‑‑1200 /um, continuous with stem or at an angle to it, base with rhizoids.  Sporophyte capsule globose, brown; elaters 2‑spiral, brown. Spores 15‑‑18 /um, finely granulate, brown. 


Peaty soil or rock along streams; endemic, southern Appalachian Mountains of Ga., Ky., N.C., S.C., Va., W.Va.; Asia (East).


4.  Nardia geoscyphus (De Notaris) Lindberg, Brit. Hep. 27. 1875


  Alicularia geoscyphus De Notaris, Mem. Acad. Torino Ser. 2. 18: 486. 1859


Plants with shoots 5‑‑10 x 0.8‑‑1.3 mm, prostrate with ascending tips, in small flat patches or mats of suberect plants, green to brown or reddish‑brown, often purplish beneath.  Stems 275‑‑325 /um in diameter; branches few, intercalary; rhizoids dense, scattered along stem, colorless, occasionally reddish tinged.  Leaves distant to contiguous on lower stem to imbricate on distal stem, slightly concave, orbicular to reniform, 450‑‑575   x 750‑‑900 /um, entire to shallowly retuse, or 2-lobed with sinus less than 1/5 leaf length, forming blunt, rounded, entire lobes; median leaf cells 24‑‑30  x 20‑‑25 /um, marginal cells smaller, 18‑‑25 /um; cuticle smooth; cell walls thin, trigones large to bulging; oil bodies 2‑‑3 per cell, ovoid to ellipsoid, large, 7‑‑15  x 6‑‑10 /um, granular‑opaque.  Underleaves vestigial to subulate or lanceolate, often connate to leaf on one side, largest near stem apex.  Sexual condition paroicous.  Androecia beneath gynoecia; bracts 2‑‑4 pairs, similar to leaves, larger, concave, entire to emarginate or crispate.  Gynoecia terminal, fleshy; bracts larger and broader than leaves, reniform, ca. 700  x 1000 /um, exceeding the perianth, shallowly 2--3-lobed; bracteole large, to 650 /um, sometimes lobed; perianth conical, 250‑‑300 /um, shorter than bracts, mouth crenulate‑denticulate; perigynium fleshy, 500‑‑800 /um, densely rhizoidous, continuous with upright stem or at distinct angle to prostrate stem.  Sporophyte capsule subglobose, brown; elaters 2‑spiral. Spores 14‑‑16 /um, slightly verruculose.  


Thin soil over rock outcrops or on damp peaty soil along streams, Arctic-alpine; Greenland; Alta., B.C., Nfld, N.S., Que.; Alaska, Colo., Conn., Calif., Maine, Mass., Mont., N.H., N.J., Ohio, Oreg., Pa., Utah, Wash., Wyo.; Europe; Asia.


A varietal name, Nardia geoscyphus var. bifida R. M. Schuster, has been proposed for plants with all leaves emarginate or shallowly 2-lobed with rounded lobes and decurrent leaf bases.  This variety is known from specimens collected in northeastern Greenland from soil in rock caves.  Variation in leaf shape may have been induced by unique environmental conditions.


5.  Nardia insecta Lindberg, Musci Scand. 8. 1879


    Nardia geoscyphus var. insecta (Lindberg) L. Clark & Frye, Bryologist 40: 15. 1937


Plants with shoots 10‑‑30 x 1.2‑‑1.8 mm, prostrate with ascending apices, light green with reddish lower stem and leaf bases.  Stems soft, ca. 300 /um in diameter, branches few, intercalary; rhizoids numerous, dense at leaf bases with few scattered along stems, colorless, occasionally slightly purplish tinged.  Leaves somewhat quadrate, wider than long, 0.6‑‑0.8 x 0.8‑‑1 mm, uniformly emarginate to 2-lobed up to 1/3 the leaf length, with triangular, blunt lobes; median leaf cells 35‑‑40  x 32‑‑36 /um, marginal cells smaller, 30‑‑33 /um; cuticle smooth, walls thin with bulging trigones; oil bodies 2‑‑3 per cell, ovoid to ellipsoid, 6‑‑7  x 14‑‑16 /um, grayish‑opaque.  Underleaves present throughout, spreading, lanceolate with reddish bases.  Sexual condition paroicous.  Androecia beneath perianth; bracts undulate‑crispate, ca. 1‑-1/2 times larger than leaves, 2-lobed ca. 1/2; base slightly concave.  Gynoecia form a terminal fleshy head, continuous with stem in upright plants, forming an angle with stem in prostrate plants; bracts wide, 700‑‑900  x 1100‑‑1400 /um, 2-lobed about 1/2 their length, crispate; bracteole lanceolate, large, to 900 /um; perianth short, 400‑‑500 /um, conical, contracted to crenulate mouth; perigynium fleshy, 1000‑‑1200 /um, elaters  2‑spiral. Spores 20‑‑24 /um, slightly granulate, brownish. 


Moist to wet humus or loam in bogs or along streams, Arctic-alpine; B.C., Nfld., N.S.; Maine, N.H., N.Y., Wash., Wyo.; Europe; Asia.


Nardia insecta is similar to and perhaps derived from N. geoscyphus and was at one time considered a form or variety of the latter.  The chromosome number of N. geoscyphus is n=18; it is n=36 in N. insecta, a slightly more robust plant of similar habitat.  Leaves of N. insecta are almost all 2-lobed to about 1/3 their length and cells are slightly larger with coarser trigones.


6.  Nardia breidleri (Limpricht) Lindberg, Meddel. Soc. F. et Fl. Fennica 6: 252. 1881


    Alicularia breidleri Limpricht, Jahresb. Schles. Gesell. Vaterl. Kult. 57: 311. 1880


Plants minute, with shoots 1‑‑4 x 0.3‑‑0.5 mm, in small patches, light green to reddish‑brown or purplish, with numerous ventral stolon‑like branches bearing small leaves.  Stems soft, 100‑‑150 /um in diameter; branching ventral or lateral intercalary; rhizoids scattered along ventral stem, colorless.  Leaves remote to contiguous, orbicular to oblong, slightly concave, 165‑‑325  x 160‑‑275 /um, entire to retuse or 2-lobed to 1/4, the lobes unequal with the dorsal smaller, lobe apices rounded, sinus obtuse; median leaf cells 15‑‑24  x 14‑‑16 /um, marginal cells smaller, 12‑‑14 /um; cuticle smooth; walls slightly thickened, trigones small or absent; oil bodies 1‑‑3 per cell, small, 3‑‑10 /um, homogeneous.  Underleaves subulate, occasionally with a lateral tooth, apparent only at stem apex.  Sexual condition dioicous.  Androecia terminal, becoming intercalary; bracts imbricate to julaceous, in 4‑‑7 pairs, concave, 2-lobed, occasionally with a lateral tooth, wider than long, larger than leaves, ca. 250  x 270 /um, purplish; antheridia 1‑‑2 per bract, stalk 2‑seriate.  Gynoecia on thick main stem or short branch with a fleshy rhizoid-berring perigynium at right angle to stem; bracts orbicular to reniform, concave; bracteole oblong to lanceolate; perianth conical, short, ca. 300 /um, hidden by bracts, mouth crenulate; perigynium fleshy, ca. 800 /um.  Sporophyte capsule globose, brown; elaters 3‑‑4 spiral. Spores 9‑‑12 /um, slightly granulate.


Wet soil in snow melt, deep water of lake, Arctic-alpine; w Greenland; Alta., B.C., Oreg., Wash.; Europe; Asia.


SELECTED REFERENCE  Wagner, D.H, J. Christy, and D. Larson. 2000.  Deep-water bryophytes from Waldo Lake, Oregon. Lake and Reservoir  Management.  16: 91--99


7.  Nardia assamica (Mitt.) Amakawa, J. Hattori Bot. Lab. 25: 23. 1963


Jungermannia assamica Mitt. J. Proc. Linn. Soc., Bot. 5: 91. 1860 [1861]


Plants 20‑‑70 x 0.4-0.7 mm, prostrate to ascending, forming loose mats, whitish and whitish green to brown.  Stems fleshy, 50‑‑100 /um in diameter, branches few, intercalary; rhizoids few to numerous, colorless, often absent near stem apex.  Leaves imbricate to distant, oblique- to erect-spreading and obliquely oriented or slightly laterally appressed, orbicular to reniform, ca. 0.37‑‑0.45 x 0.45‑‑0.5 mm, with rounded, entire apices or shallowly retuse or truncate apex, concave, barely decurrent dorsally; median leaf cells 30‑‑38  x 25‑‑30 /um, marginal cells smaller, subquadrate, 20‑‑27 /um; cuticle smooth; trigones distinct, small; oil bodies 1 per cell, present in ca. 30% of leaf cells, spheric to shortly ellipsoidal, 10‑‑18  x 10--16 /um, brownish, coarsely granulate.  Underleaves spreading, spathulate, relatively large, 0.2--0.25 x 0.19--0.22 mm. Sexual condition dioicous.  Androecia terminal, becoming intercalary; bracts in 3‑‑8 pairs, similar to leaves.  Gynoecia terminal on main stems, bracts inserted on perigynium, larger and broader than leaves, reniform, hardly exceeding length of perianth, hiding it or perianth shortly exerted; bracteole spathulate, commonly connate to bracts; perianth conical, colorless to purplish, mouth crenulate; perigynium swollen, often purplish, continuous with stem, ca. 2--3 times longer than perianth.  Sporophyte capsule brown; elaters 2‑spiral. Spores 14‑‑15 /um, slightly papillose, brown.


Wet exposed soil with water seeping, broadly East Asian-North Pacific; Alaska; Asia.


Nardia assamica is characterized by creeping growth, distant leaves, spathulate underleaves commonly connate with the leaves, and coarsely granulate brownish oil bodies, 1 per cell, present in ca. 30% or less of leaf cells.


8.  Nardia hiroshii Amakawa, J. Hattori Bot. Lab. 21: 283, fig. 9: m--v. 1959


Plants with shoots 5‑‑10 x 1.1‑‑1.4 mm, ascending to (rarely) creeping, whitish to pale green and yellowish, commonly with apical portions of the leaves brownish golden.  Stems soft, 180--300 /um in diameter, branches few, intercalary; rhizoids numerous, dense at underleaf bases with few scattered along stems, colorless to brownish.  Leaves contiguous to distant, subtransversely to subobliquely inserted (at ca. 30-45° with stem axis), with barely or shortly decurrent dorsal leaf base; divided by obtuse-angular to loosely gibbous sinus descending for 1/5--1/4 of leaf length into two triangular obtusely apiculate lobes, mostly deeper brownish golden colored near lobe apices; moderately concave to concave-canaliculate, transversely elliptic to trapezoidal, 0.65--0.75 x 0.75--0.85(--0.9) mm (length to width ratio is 1:0.85--0.95); midleaf cells mostly 5--6-gonal, 28--53 x 25--42 /um, thin-walled, trigones moderate in size, triangular to convex, walls colorless to pale yellowish; near lobe apex walls brownish, trigones triangular to convex, lumen rounded, ca. 28--34 μm in diameter; near the base 30--47 x 25--41 /um, thin-walled, trigones convex, walls colorless; oil-bodies in the midleaf cells 2--5 per cell, 11--20 x 8 /um, irregularly elliptic, finely granulate.  Underleaves present throughout, laciniate to narrowly triangular, rarely with additional unicellular tooth near base, sometimes connate with ventral base of one leaf of each pair; rarely underleaves hidden in the rhizoids and then invisible, 3--6 cells wide at the base and 5--8(--10) cells long (ca. 200--280 x 60--140 /um).  Sexual condition dioicous.  Androecia intercalary, 2-lobed ca. 1/2; base slightly concave.  Gynoecia terminal; perianth conical, loosely plicate, ca. 0.5 x 0.8 mm, with loosely beaked mouth, hidden within bracts; perigynium not pendent, strongly rhizogenous, ca. 1.5 mm; bracts similar to leaves, but more deeply lobate and having obviously apiculate lobes, undulate and crispate at margin; bracteoles become bigger to the pair adjacent to the perianth, vary from 0.4 mm in third pair (downward from the perianth) to 0.8 mm in upper pair, triangular to narrowly triangular, the biggest bracteole undulate at margin. Sporophyte capsule brown; elaters 2‑spiral. Spores 13‑‑15 /um, slightly granulate, brownish. 


Soil at edge of meadow; bedrock granite, boreal, mainly East Asian (only one record in western hemisphere); Calif.; e Asia.


9. Nardia japonica Stephani, Bull. Herb. Boissier 5: 101. 1897


Plants 10--20 x  0.5--0.8(--1.2) mm, green to golden-brown or reddish.  Stems thick, 160--208 /um in diameter, branches rare; rhizoids common, colorless, in obliquely spreading fascicles from ventral side of stem mostly near underleaf base. Leaves contiguous to distant, slightly decurrent at the dorsal insertion, trapezoidal to ovate, distinctly 2-lobed, concave to almost plane, 0.2--0.3 x 0.3--0.4 mm, lobes broadly triangular; lobe apices obtuse to acute; sinus descending 1/4--1/5 leaf  length, sinus base rounded to obtuse; median cells 18--30(--32)  x 22--40 /um; marginal cells 12--24 x (16--)20--34 /um; cell walls becoming golden to brownish, thin with bulging trigones; oil-bodies homogeneous, shining, occasionally becoming segmented; 2--4 per cell, nearly spherical, 3--9 /um in diameter, or ovoid-ellipsoid and 4--8 x 6--17.5 /um. Underleaves spathulate to triangular-lanceolate, large, distributed along entire length of stem, 0.24--0.4 x 0.1r--0.23 mm. Sexual condition dioicous. Androecia intercalary, with 4–6 pairs of bracts, nearly spicate. Gynoecia terminal; bracts in 2--3 pairs, similar to but larger than leaves; bracteoles similar to but larger than underleaves; perigynium short-pendent, fleshy,  larger toward the ventral side of stem. Sporophyte capsule brown; elaters 2-spiral. Spores 14.5--23 /um, finely papillose, red-brown.


Shaded, moist soil, humus, gravel, open subalpine meadows, near stream, ; 300--1900 m; B.C.; Alaska, Oreg., Wash.; Asia (China, Japan, Russia in e Siberia).


Nardia japonica is rather small compared to other species in the genus, but abundant when found. Distinguishing characters are 2-lobed leaves, large underleaves along the length of the stem, and dioicous.


SELECTED REFERENCES  Bakalin, V. A. and K. G. Klimova. 2016. A note on Nardia japonica Steph. (Gymnomitriaceae). Bot. Pacifica 5: 43--50.  Godfrey, J. D. and G. A. Godfrey. 1980. Notes on hepatics in the Pacific Northwest. Bryologist 83: 224--228.


4. PRASANTHUS Lindberg in Kgl. Sv. Vetensk. Akad. Handl. 23 (5): 62. 1989 * [Greek prason, onion, plus anthos, flower, alluding to the fleshy perigynium]

Won Shic Hong