BFNA Title: Nardia
XXX. 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 \mu, sometimes reddish tinged, not distinctly differentiated from slightly longer medullary cells, 35‑‑60 x 16‑‑40 \mu; 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, 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 \mu, 2‑‑4 spiral. Spores 9‑‑24 \mu.
Species 14 (8 in the flora), moist soil or humus: North America, South America (Brazil), Europe, Asia, Africa.
The genus is distinguished by the wide, entire to 2-lobed leaves, the lanceolate 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 more than 0.5 mm wide; leaves entire or 2-lobed; trigones distinct in leaf cells.
2. Plants paroicous, androecia beneath perianth.
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.
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 of leaf length, lobes acute; oil bodies 2--5 per midleaf cell; underleaves laciniate to narrowly triangular … 8. Nardia hiroshii
4. Leaves unlobed.
6..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
6. 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 .
7. Leaves orbicular, marginal leaf cells only slightly smaller than median cells; stem hyalodermis absent ......... 1. Nardia scalaris
7. 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 \mu, 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 \mu, 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, granular botryoidal, made up of numerous 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 \mu, marginal cells smaller, 20‑‑30 \mu, oil bodies 2‑‑3 per cell, ovoid to ellipsoid, 8‑‑15 x 6‑‑7 \mu, 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 \mu, marginal cells 28‑‑38 \mu, oil bodies 2‑‑3(‑‑6) per cell, ovoid to ellipsoid, 10‑‑16 x 6‑‑9 \mu, 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 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 \mu 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 \mu, marginal cells smaller, subquadrate, 18‑‑25 \mu; cuticle smooth; trigones distinct, small to large and bulging; oil bodies 1‑‑3 per cell, ovoid to ellipsoid, 10‑‑14 x 7‑‑10 \mu, 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 \mu, 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 \mu 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 \mu, marginal cells smaller (20‑‑24 \mu); cuticle smooth to slightly verruculose, walls thin, trigones large, bulging; oil bodies 3‑‑5 per cell, ovoid to ellipsoid, 7‑‑10 x 10‑‑16 \mu, 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 \mu, mouth entire or irregularly lobed and crenulate; perigynium fleshy, as long as perianth or longer, 700‑‑1200 \mu, continuous with stem or at an angle to it, base with rhizoids. Sporophyte capsule globose, brown; elaters 2‑spiral, brown. Spores 15‑‑18 \mu, 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 \mu 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 \mu, 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 \mu, marginal cells smaller, 18‑‑25 \mu; cuticle smooth; cell walls thin, trigones large to bulging; oil bodies 2‑‑3 per cell, ovoid to ellipsoid, large, 7‑‑15 x 6‑‑10 \mu, 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 \mu, exceeding the perianth, shallowly 2--3-lobed; bracteole large, to 650 \mu, sometimes lobed; perianth conical, 250‑‑300 \mu, shorter than bracts, mouth crenulate‑denticulate; perigynium fleshy, 500‑‑800 \mu, densely rhizoidous, continuous with upright stem or at distinct angle to prostrate stem. Sporophyte capsule subglobose, brown; elaters 2‑spiral. Spores 14‑‑16 \mu, 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 only 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 \mu 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 \mu, marginal cells smaller, 30‑‑33 \mu; cuticle smooth, walls thin with bulging trigones; oil bodies 2‑‑3 per cell, ovoid to ellipsoid, 6‑‑7 x 14‑‑16 \mu, 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 \mu, 2-lobed about 1/2 their length, crispate; bracteole lanceolate, large, to 900 \mu; perianth short, 400‑‑500 \mu, conical, contracted to crenulate mouth; perigynium fleshy, 1000‑‑1200 \mu, elaters 2‑spiral. Spores 20‑‑24 \mu, 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 \mu 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 \mu, 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 \mu, marginal cells smaller, 12‑‑14 \mu; cuticle smooth; walls slightly thickened, trigones small or absent; oil bodies 1‑‑3 per cell, small, 3‑‑10 \mu, 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 \mu, 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 \mu, hidden by bracts, mouth crenulate; perigynium fleshy, ca. 800 \mu. Sporophyte capsule globose, brown; elaters 3‑‑4 spiral. Spores 9‑‑12 \mu, slightly granulate.
Wet soil in snow melt, Arctic-alpine; w Greenland; Alta., B.C., Wash.; Europe; Asia.
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 \mu 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 \mu, marginal cells smaller, subquadrate, 20‑‑27 \mu; 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 \mu, 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 \mu, 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 \mu 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 \mu, 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 \mu, 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 \mu). 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 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 \mu, slightly granulate, brownish.
Soil at edge of meadow; bedrock granite, boreal, mainly East Asian (only one record in western hemisphere); Calif; e Asia; Europe (introduced).