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BFNA Title: Nardia |
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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 [1861] Plants 20‑‑70 x 0.4-0.7 mm, prostrate to
ascending, forming loose mats, whitish and whitish green to brown. Stems
fleshy, 50‑‑100 \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). |
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