XX. GYMNOMITRIACEAE H. Klinggraff
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
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
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--
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--
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,
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--
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--
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--
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--
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--
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.
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 (
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
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
8. Plants very small with sterile shoots not
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
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
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--
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.
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
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
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
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.
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--
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--
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
Montane species, on moist
soil-covered rocks in mountains; 1700--
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--
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--
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--
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
Montane pioneer on damp acidic rocks, on banks of
stony streams, rarely in standing water; medium to high elevation (
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--
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--
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
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
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 
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