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BFNA Title: Lepidoziaceae |
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XX. LEPIDOZIACEAE
Limpricht Vadim Bakalin Plants 1--3-pinnate or pseudodichotomous, flagelliform branches frequent, rhizoids
rare, originated near underleaf bases. Leaves
incubous to transverse, 3--4(--5)-lobed, leaf cells with several oil bodies or
oil bodies as numerous minute droplets or absent. Underleaves always regular, lobed, rarely
nearly entire. Sexual condition dioicous
or autoicous. Gynoecia on short isophyllous postical-intercalary branches. Perianth conical to fusiform, more or
less distinctly 3-plicate in its distal half. Capsule ovoid to ovoid-cylindric with 2--6-stratose wall. Specialized asexual reproduction by caducous
leaves. SELECTED REFERENCES
Frey W. 2009. Syllabus of Plant Families. Adolf Engler's Syllabus
der Pflanzenfamilien. Part 3: Bryophytes and Seedless Vascular Plants. 13th ed.
Fulford, M. 1936. The genus Bazzania in the United States and Canada. Amer. Midland Nat. 17: 385--424.
Schuster R. M. 1969. The Hepaticae and Anthocerotae of North America. Vol. 2.
New York, Columbia University Press. Hong,
W. S. 1988. The family Lepidoziaceae in North America
west of the hundredth meridian. Bryologist 91:326--333. 1. Leaves shortly
incised or lobed less than 1/5 of leaf length, plants commonly larger than 2 mm
wide … 1. Bazzania, p. XXX 1. Leaves divided
more than 1/3 of leaf length, plants commonly narrower than 2 mm wide. 2. Leaves divided
to 1/3--2/3 of leaf length, plants wider than (0.4--)0.5
mm… 3. Lepidozia, p. XXX 2. Leaves divided
almost to the base (with undivided portion 1--2 cells high), plants narrower than
0.45 mm. 3. Hyaloderm absent (North American taxa), branches of Frullania and Microlepidozia types,
oil bodies virtually absent, or as minute droplets, cells to 30--35 \um long…
2. Kurzia, p. XXX 3. Hyaloderm well-defined, branches of Frullania type only, oil bodies
distinct, leaf cuticle smooth, cells to 90--110 \um long in the lobe middle …
4. Telaranea, p. XXX 1. BAZZANIA Gray Nat.
Arr. Brit. Pl. 1: 704, 1821 * [For M. Bazzani, Italian
professor of anatomy] Dendrobazzania R.M. Schust. & W.B. Schofield Plants prostrate to ascending,
rarely erect, in dense to loose patches, deep green to greenish yellowish and
brownish, rarer reddish, rusty to purple-brown, greatly varying in size (from
0.7 to 5.5 mm in regional taxa). Stem
freely pseudodichotomously terminally (Frullania--type)
branched for normal branches, and also commonly with numerous ventral intercalary
flagellae. Rhizoids virtually absent
or scarce, present as short erect-spreading fascicles originating near ventral
underleaf base. Leaves incubous, 1--4-lobed,
sometimes with additional teeth along margin (mostly confined to apical third
of the leaf), obliquely inserted, slightly to strongly
convex, in some taxa strongly curved to ventral side, easily deciduous in some
taxa. Underleaves appressed to the
stem or obliquely to erect-spreading, plane to recurved
along margin, with apex commonly lobed or coarsely crenulate and dentate, rarely
almost entire, chlorophyllose or hyaline along margin. Midleaf cells mostly pachydermous, rarely
leptodermous; cuticle smooth to (rarely) papillose;
oil bodies mostly smooth surfaced, botryoidal, rarer homogenous to homogenous
biconcentric, or ellipsoidal and having finely granulate
surface. Sexual condition dioicous (but commonly sterile), with archegonia
and androecia appearing on short ventral intercalary branches. Perianth long--conical to fusiform, commonly
with three main plicae; perigynium absent. Female bracts ovate, similar in shape and size to bracteoles, imbricate.
Androecia male bracts spicate, in 4--6
pairs, ovate, cupped, entire to 2-lobed, 1--2-androus. Capsule ovoid to oblong-ovoid, capsule wall 4--6-stratose; with robust
seta of ca. 16 epidermal and numerous internal rows. Elaters 2-spiral. Spores
small, brown. Species 100--150 worldwide,
everywhere except Antarctic, 7 in the flora. SELECTED REFERENCES Fulford, M. 1936. The genus Bazzania in the United States and Canada. Amer. Midland Nat. 17: 385--424.
Schuster, R.M. 1969. The Hepaticae and Anthocerotae of North America. Vol. 2.
New York, Columbia University Press. Hong,W. S. 1988. The family Lepidoziaceae in North America west of the hundredth meridian.
The Bryologist 91:326--333. 1. Underleaves transversely
elliptic to shallowly retuse, trigones large and convex at midleaf … 5. Bazzania pearssonii 1. Underleaves distinctly
lobed, incised or at least toothed, trigones large, convex
to small and concave. 2. Plants brown to rusty colored (at least with admixture of
these colors). 3. Leaves freely caducous, with apex 2--3-lobed … 4. Bazzania nudicaulis 3. Leaves not caducous, with apex 3-lobed. 4. Trigones in midleaf cells large, convex, leaf apex ca. 1/2
of basal part in width, with slightly diverging lobes [rare meta-Himalayan taxon
recorded in Pacific North-West] … 3. Bazzania griffithiana 4. Trigones in midleaf cells moderate in size to small, concave,
leaf apex 1/4--1/3 of basal part in width, lobes straight or convergent [widely
distributed circumpolar taxon] … 6. Bazzania tricrenata 2. Plants green to salad- or deep green, never brown or rusty. 5. Plants 3--6 mm wide, underleaves distinctly dentate or lobed
with acute lobe apices … 7. Bazzania trilobata 5. Plants less 2 mm wide, underleaves lobed (rarely lobed),
with margin entire and lobe apices rounded. 6. Underleaves as long as wide, with middle sinus deeper than
others (if present), leaves frequently 2-lobed at the tips, oil bodies 2--4 per
midleaf cell … 1. Bazzania ambigua 6. Underleaves wider than long, with middle sinus similar in
size with other, well-developed leaves mostly 3-lobed, oil bodies 8--12 per midleaf
cell … 2. Bazzania denudata 1. Bazzania ambigua (Lindenberg) Trevisan,
Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat.
4 (13): 414, 1877 E Mastigobryum ambiguum Lindenberg, Syn. Hepat., 217. 1845 Plants brownish green to
brown, 15--40 mm long, and 1000--1800 \um wide, more or less rigid, forming loose
patches. Stem 200--450 \um in diameter,
easily branched, with numerous or a few (forms with caducous leaves) ventral flagellae.
Rhizoids few, in flagelliform branches
and the bases of sexual branches. Leaves
contiguous to
imbricate, convex, commonly curved to dorsal side, but not inrolled, dorsally insertion line oblique to subtransverse
or loosely arcuate, ventrally leaf connate with underleaf or not connate, loosely
falcate, obliquely ovate and oblong-ovate, mostly shallowly 2-lobed, rarely unlobed
or 3-lobed (2-dentate leaves always dominant), with lobes more or less acute,
leaf margin entire to loosely crispate, 700--900 x 450--600 \um. Underleaves connate with one leaf of the
pair or not connate, quadrate-orbicular, usually squarrose, 300--450 \um in diameter,
2--4-lobed (rarely unlobed), with rounded lobes, if 3-lobed, the middle sinus
commonly deeper than marginal. Midleaf
cells ca. 20 \um in diameter, thin-walled, trigones small, but distinct, concave
to triangle, cuticle smooth; lobe apices with smooth cuticle; oil bodies 2--4(5)
per cell, spherical to fusiform, with amorphous texture, neither homogenous
nor glistening. Sexual condition dioicous. Perianth narrowly ovate, ca 3 mm long and 1 mm wide bracts and bracteole
similar, 1000--1300 x 800--1200 \um, with slime papillae along margin, with short
teeth, not divided by conspicuous sinus. Androecia
branches 0.5 mm long, bracts in 4--6 pairs, 1-androus. Capsule 1000 x 500 \um. Elaters
200--250 x 8--10 \um. Spores brownish,
12--14 \um. Acidophilic mesophyte, tree trunks and logs, rarely on rocks (then produces
rusty-brown colored modifications); elevation 0--500 m; B.C.; Alaska, Idaho, Oreg.,
Wash. Bazzania ambigua is frequently associated with Blepharostoma trichophyllum,
Calypogeia
spp., Cephalozia bicuspidata, Fuscocephaloziopsis lunulifolia,
Lepidozia reptans, Mylia taylorii, etc.
2. Bazzania denudata (Torrey ex Gottsche, Lindenberg & Nees) Trevisan, Mem. Reale Ist. Lombardo
Sci., Ser. 3, Cl. Sci. Mat. 4: 414, 1877 Mastigobryum denudatum Torrey ex Gottsche, Lindenberg & Nees, Syn. Hepat. 2: 216, 1845; Bazzania denudata subsp. ovifolia (Stephani) S. Hattori;
B. ovifolia
(Stephani) S. Hattori Plants deep green to brownish
green, yellowish green and brownish green, 15--40 mm long and 1000--2500 \um wide,
more or less rigid, forming loose patches. Stem 200--450 \um in diameter, easily branched, with numerous or a
few (forms with caducous leaves) ventral flagellae. Rhizoids virtually absent, rarely a few, in erect fascicles originating
near underleaf bases. Leaves mostly
contiguous, but also subimbricate or distant, not or
easily deciduous, convex, turned to dorsal side when wet, dorsal insertion line
oblique to subtransverse or loosely arcuate, ventrally leaf connate with underleaf
or not connate, obliquely ovate, distinctly 3-lobed with lobes more or less acute
lobes or (1--)2--3-lobed with lobes obtuse, leaf margin
entire to loosely crispate, 700--1250 x 500--880 \um. Underleaves connate with both or with one leaf of the pair or (rarer)
not connate, transversely elliptic, erect-spreading, plane or recurved, 3--5-lobed
or crispate, lobes (if present) obtuse to round at apices. Midleaf cells 20--32 x 20--27 \um, thin-walled,
trigones moderate in size, convex, cuticle smooth; lobe apices with smooth cuticle;
cells along leaf margin 20--25 \um, more or less thin--walled, with large, slightly
convex trigones, external wall noticeable to moderately thickened, cuticle smooth;
oil bodies 8--12 per cell, homogenous, glistening, spherical to oblong and shortly
fusiform, occasionally segmented. Sexual condition dioicous. Perianth unknown, female bracts with rounded
or shortly 2-lobed apex. Androecia
unknown. Capsule unknown. Elaters unknown. Spores unknown. Decaying wood, humus
on steep slopes, cliff crevices, avoids dry substrata, prefers merely shaded habitats,
rarely occurring in open sites or in full shade; elevation 0--2000 m; Alta., B.C.,
Nfld. and Labr., N.S., Ont., Que.; Alaska, Calif., Conn., Ga., Ky., Maine, Mass.,
N.Y., N.C., Ohio, Oreg., Pa., S.C., Tenn., Vt., Va., Wash.; Eurasia. An acidophilic mesophyte, Bazzania denudata occupies many types of habitats. The species
is relatively more plastic ecologically and sometimes occurs with other taxa of
the genus. 3. Bazzania griffithiana
(Stephani) Mizutani, Journal Hattori Bot. Lab. 30: 82,
1967 Mastigobryum griffithianum Stephani, Bulletin de l'Herbier Boissier, sér. 2 8: 959, 1908; Dendrobazzania griffithiana
(Stephani) R.M. Schuster & W.B. Schofield Plants yellowish or brownish
to golden brown, never red, forming loose patches, 20--80 x mm long and 1000--1500
\um wide. Stem easily pinnately branched,
ca. 400 \um in diameter, ventral flagellae rarely present. Rhizoids virtually absent. Leaves not caducous, contiguous to distant
or subimbricate, obliquely ovate--triangular, with obliquely
truncate leaf apex, dorsal insertion line subtransverse to arcuate, ventrally
not connate with leaves or connate with one leaf of the pair, 1000--1500 x 1200--1500
\um, strongly turned to ventral side, convex, 3-lobed, lobes subequal to unequal, acute, slightly diverging, sinus V-shaped.
Underleaves erect-spreading, not connate
or connate in one side with corresponding leaf, rounded, 3--4-lobed, with rounded
lobes, strongly auriculate at the base. Midleaf
cells pachydermous, with large convex to bulging trigones,
20--25 x 20--23 \um, somewhat larger basally and smaller apically; oil bodies
mostly smooth surfaced or faintly septated, ovoid to
ellipsoidal, in the midleaf (1--2--)3--5(--7) per cell, ca. 4--5.5 x 2.5--4 \um.
Sexual condition unknown, probably dioicous. Perianth unknown, female bracts divided near apex into 2--3 lacinae by narrow sinus. Androecia unknown. Capsule
unknown. Elaters unknown. Spores unknown. Generally acidophilic
mesophyte, maritime forests; elevation low, below 500m;
B.C.; Eurasia. Bazzania griffithiana prefers mesic open to partly shaded cliff crevices and open
(but not in full sun) rocks, rarely occurring in shady sites. 4. Bazzania nudicaulis A. Evans, Bryologist 26: 62. 1923 [1924] E Plants yellowish brown to
reddish brown, 8--30 mm long, 750--1600 \um wide, more or less rigid, forming
loose patches. Stem 140--230 \um in
diameter, sparsely branched, branches distant, widely diverging, ventral flagellae
few. Rhizoids virtually absent, rarely
a few, in erect fascicles originating near underleaf bases or on bases of leaves
of flagellae. Leaves distant to subimbricate, commonly easily deciduous, slightly to moderately
convex, turned to dorsal side when wet and dry, dorsal insertion line oblique
or loosely arcuate, ventrally leaf connate with underleaf or not connate, ovate
to ovate-lanceolate, nearly symmetrical, distinctly 2- to 3-lobed with lobes more
or less acute, leaf margin entire to loosely crispate, 700--1250 x 500--880 \um.
Underleaves loosely connate with both
or with one leaf of the pair or not connate, suborbicular or quadrate-orbicular,
erect-spreading, plane to slightly convex, 2--3-lobed or crispate, lobes (if present)
obtuse to round at apices. Midleaf
cells 20--23 x 20--31 \um, thin-walled, trigones moderate in size, convex, cuticle
smooth to loosely verrucose; cells along leaf margin 20--25 \um, more or less
thin--walled, with small, but distinct, concave trigones, external wall moderately
thickened, cuticle smooth; oil bodies 1--4 per cell, finely granulate, spherical
to ovoid and ellipsoidal. Sexual condition and else unknown. Acidophilic mesophyte occurring mostly on moist to mesic rocks and rarer
on bark of trees; elevation 2500--3500 m; N.C., Tenn., Va. Bazzania nudicaulis occurs on the “fog-shrouded
summits of the highest peaks of the Appalachian system” (R. M. Schuster, 1969:
81). This species is commonly associated with Herbertus, Frullania and Plagiochila. In
rocks sometimes admixed with closely allied Bazzania tricrenata. Prefers partly shaded habitats
although sometimes growing in relatively well exposed sites. 5. Bazzania pearsonii Stephani, Hedwigia 32 (4): 212. 1893 Plants greenish to golden-green,
pale brown and reddish-brown, 40--80 mm long, 1100--1400 \um wide, more or less
rigid, forming loose patches. Stem
250--300 \um in diameter, easily branched, somewhat circinate at apex. Rhizoids virtually absent, rarely a few,
in erect fascicles originating near underleaf bases. Leaves mostly contiguous to imbricate, not caducous, convex, turned
to dorsal side when wet and dry, dorsally insertion line oblique to subtransverse
or loosely arcuate, ventrally leaf connate with underleaf or not connate, obliquely
triangular-ovate, distinctly 2--3-lobed with more or less acute lobes, leaf margin
entire to loosely crispate, 900--1200 x 700--850 \um. Underleaves spreading, with margin slightly recurved, nearly transversely
elliptic, apex entire to emarginate. Midleaf
cells with large, bulging trigones (then lumen sometimes stellate), cuticle smooth
to striolate, 22--26 \um in diameter. Sexual condition dioicous. Perianth unknown, archegonia on short
lateral branches. Androecia with 8--10
pairs of bracts. Capsule unknown. Elaters unknown. Spores unknown. Acidophilic meso-hygrophyte, peaty humus on hummocks, among Sphagnum and other bryophytes, also acidic
rocks and tree base; elevation unknown, probably below 500 m; B.C.; Alaska; Eurasia. 6. Bazzania tricrenata (Wahlenberg) Trevisan,
Mem. Reale Ist. Lombardo Sci., Ser. 3, Cl. Sci. Mat.
4: 415, 1877 E Jungermannia tricrenata Wahlenberg, Fl. Carpat. Princ.: 364, 1814; Bazzania tricrenata var.
fulfordiae
W. S. Hong Plants yellowish brownish
to greenish brown, forming loose patches, 20--40 mm long, 750--1250 \um wide.
Stem easily branched, 125--225 \um
in diameter, ventral flagellae present, but not numerous. Rhizoids virtually absent, rarely present
at the ends of ventral branches. Leaves
not caducous, contiguous to distant or subimbricate,
obliquely ovate--falcate, with attenuate leaf apex, dorsal insertion line subtransverse
to arcuate, ventrally not connate with leaves or connate with one leaf of the
pair, 750--1000 x 500--750 \um, strongly turned to ventral side, convex, (2--)3-lobed,
lobes strongly unequal, acute, sinus U--to V-shaped. Underleaves erect-spreading, not connate or connate in one side with
corresponding leaf, not recurved, 3--4-lobed, with rounded lobes, transversely
elliptic. Midleaf cells 17--23 x 17--20
\um, thin--walled, trigones moderate in size, concave to triangular, cuticle smooth;
cells along margin 12--20 \um, external wall not or loosely thickened, cuticle
smooth; cuticle in lobe apices smooth; oil bodies 2--4(--5) per cell, smooth surfaced,
glistening, shortly oblong to nearly spherical or fusiform, occasionally biconcentric (having one pupil). Sexual condition dioicous.
Perianth ovoid-cylindrical, female
bracts divided near apex into 2--3 lacinia by narrow sinus. Androecia spicate, with 1-several pairs
of bracts. Capsule ellipsoidal; seta
1--2 cm, with 16 outer and 20--50 inner cells. Elaters 2-spiral, 260--500 x 6--8 \um. Spores 15--20 \um, papillose. Generally neutrophilic,
although acido- and basic-tolerant, mesophyte, mesic cliff crevices, open (but not in full sun)
rocks, rarely in shady sites, prefers rocky substrates, sometimes well-exposed
humus on steep slopes; elevation 0--2300 m; B.C., N.B., Nfld. and Labr. (Nfld.),
N.S., Que.; Alaska, Calif., Conn., Idaho, Ky., Maine, Mass., N.H., N.Y., Oreg.,
Pa., Tenn., Vt., Va., Wash., W.Va.; Central America; Eurasia. 7. Bazzania trilobata (L.) Gray, Nat. Arr. Brit. Pl. 1: 704, 1821 Jungermannia trilobata L., Sp. Pl. 2: 1133. 1753 Plants prostrate to erect
in more or less loose patches, deep green to (rarely) yellowish green or brownish
green, however always brightly colored), 50--170 mm long, (3--)4--6.5 mm wide.
Stem often branched, with numerous
ventral flagellae, 380--575 \um in diameter. Rhizoids virtually absent. Leaves
not caducous, convex-obcanaliculate, not turned to ventral
side when wet and mostly not turned when dry (rarely in dry condition turned to
ventral side), contiguous to imbricate or distant in weak phases, 1400--4250 x
1250--2750 \um, obliquely ovate to narrowly trapezoidal, with loosely tapered
and sometimes falcate apical 1/3, widest near base, apex (2--)3(--4)-lobed, sinus
V- to U-shaped, lobes acute,
leaves in apical (rarely also basal) third sometimes sparsely dentate or loosely
crispate along margin or margin entire, insertion line dorsally distinctly arcuate.
Underleaves appressed to the stem or
obliquely spreading, 450--900x700--1400 \um, mostly (East Asian phenotypes) 4--6-lobed,
with many additional teeth or coarsely dentate along margin and then not lobed,
commonly hyaline along margin (but cells with trigones), not connate with leaves
or connate in one side with corresponding leaf. Midleaf cells 22--45 x 17--33 \um, thin--walled, trigones moderate
to large, convex, cuticle smooth; cells along leaf margin 15--25 \um, external
wall thickened, trigones adjacent to external wall large and convex, tangential
wall thin to unequally thickened, inner wall thin, with moderate to large in size,
concave to convex trigones, cuticle smooth; cells in lobe apices with smooth cuticle;
oil bodies shortly oblong to irregular in shape, smooth surfaced to loosely segmented.
Sexual condition dioicous. Perianth
ovoid-cylindrical, 3-plicate, to 6 mm long. Androecia with 1-several pairs of bracts, monandrous.
Capsule oblong-ovoid. Elaters ca. 200 x12 \um, 2-spiral. Spores 12--15 \um, with labyrinthine
markings. Acidophilic to neutral-tolerant
mesophyte , rarely in forest over limestone, but then growing on thick
acidic litter, known only from moderately shaded habitats. elevation 0--1000
m; B.C., N.B., Nfld. and Labr., N.S., Ont., Que.; Ala., Alaska, Conn., D.C., Fla.,
Ill., Ind., Ky., Maine, Md., Mass., Mich., Minn., N.H., N.J., N.Y., N.C., Ohio,
Pa., R.I., S.C., Tenn., Vt., Va., W.Va., Wis.; Eurasia. In northern extremes
Bazzania trilobata grows
on litter between and over mosses (most commonly Pleurozium and Hylocomium) under exposed Pinus then sometimes also on overlying
branch bases) and Larix
forests. Southward it is locally abundant in mountain and coastal boreal dark
(Picea and Abies) coniferous forests, then
continues northward in mixed broadleaved-coniferous forests (hemiboreal zone or belt), where slightly rarer than in boreal
communities. It rarely descends from mixed forests to cool temperate broadleaved
forest zone, but may occur in the subtropics in Florida, where it is found in
Magnolia virginiana-Persea swamps (R. M. Schuster 1969). 2. KURZIA G.
Martens, Flora 53 (27): 417, 1870 * [For S. Kurz,
specialist in Indian flora] Microlepidozia (Spruce) Jørgensen, Bergens Mus. Skr. 16: 303, 1934 Plants prostrate to ascending,
caespitose, merely rigid, in loose patches, brownish
to green-brown, seems never red, 3--10 mm long, 200--500 \um wide. Stem freely and regularly pinnately and
2-pinnately branched (branches of Microlepidozia type) for normal branches outer cells in 8--16
rows, firm, not larger than inner cells, distinctly thick-walled, not forming
hyalodermis, vegetative branches and central stem sometimes become to microphyllous
flagellae, some flagellae also arise from axis of underleaves. Rhizoids sparse to virtually absent, except
on flagellae, as thick and short obliquely to erect spreading fascicles originating
near underleaf and reduced leaf bases. Leaves
incubous to transverse, not deciduous, imbricate to contiguous, rarely distant,
slightly convex, divided almost to the base into (3--)4(--6) subulate lobes, 1--3(--6)
cells wide, 100--300 \um long and wide, subquadrate to transversely elliptic when
flattened. Underleaves in the most
taxa similar to leaves or considerable smaller and then reduced to two subulate
and uniseriate lobes of 2--3 cells long, abaxially loosely convex. Leaf lobe cells thick-walled, oblong (1--3
as long as wide), cuticle smooth to verruculose, chlorophyllose and opaque; oil
bodies mostly absent, rarely present in some basal cells as a few to numerous
smooth surfaced spherules. Sexual condition dioicous. Perianth terminal, on short postical branches,
without vegetative leaves, 3-plicate, cylindrical to fusiform, mouth lobulated
to crenulate and ciliate, bracts in 2--3 pairs, similar to bracteoles, oblong
ovate, 2--3-lobes with sinus descending to the middle or only emarginate at the
apex. Androecia on postical branches
with reduced leaves or on elongate normal lateral branches, bracts in 2--6 pairs,
2--4-lobed to 1/2--2/3 of their length, monandrous,
antheridial stalk uniseriate, short. Capsule
wall 2-- (3)-stratose, epidermal cells sunquadrate,
with nodular thickenings, inner cells narrowly rectangular with several complete
bands of thickenings. Elaters ca. 9--12
\um thick. Spores 10--15 \um, pale
brown, verrucuose. Species ca. 30 (3
in the flora): worldwide, except Antarctic. SELECTED REFERENCES Hong,W. S. 1988. The family Lepidoziaceae
in North America west of the hundredth meridian. Bryologist 91:326--333. Schuster R. M. 1969. The Hepaticae and Anthocerotae
of North America, Vol. 2. New York, Columbia University Press. 1. Underleaves normally
4-lobed, commonly not reduced to slime papilla, heliophyte,
in peat … 2. Kurzia pauciflora 1. Underleaves normally
3-lobed, with 1- or 2-lobes commonly replaced by slime papilla, helio-sciophyte. 2. Female bract margin
short-ciliate, perianth mouth with 2--4-celled cilia, androecial bract 2-lobed
… 1. Kurzia makinoana 2. Female bract margin
obscurely dentate, perianth mouth with 1--2-celled teeth, androecial bract 3-lobed
… 3. Kurzia trichoclados 1. Kurzia makinoana (Stephani) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 171, 1963
[1964] Lepidozia makinoana Stephani, Bull. Herb. Boissier 5
(2): 94, 1897; Kurzia sylvatica (A.W. Evans) Grolle; Lepidozia sylvatica
A.W. Evans Plants dull to deep green
and brownish green, dense in low tufts,
3--10 mm long, 100--200 \um wide, creeping to loosely ascending. Stem 50--80 \um in diameter, more or less
regularly pinnately or 2-pinnately branched, with branches occasionally terminating
in flagellae. Rhizoids a few, in underleaf
bases or in flagellae. Leaves transversely
to incubously obliquely inserted, contiguous to imbricate,
subquadrate when flattened, 180--225 x 200--250 \um, merely symmetrical, deeply
3(--4)-lobed, with undivided area of 2--3 cells high, lobes entire, subulate to
narrowly lanceolate, (1--)2 cells wide in base, dorsal
lobe slightly smaller than other; branch leaves commonly smaller that stem leaves
and only 2--3-lobed. Underleaves smaller
than leaves, 3-lobed to 1--2 cells to the base, slightly convex, 115--225 \um
long and 65--120 \um wide. Leaf lobe
cells 16--26 x 14--20 \um at lobe bases, with walls slightly thickened, cuticle
almost smooth or slightly verruculose. Perianth
terminal, on short postical branches, ovoid-cylindrical to fusiform, basically
3-plicate, but become 6-plicate near mouth, mouth lobulated to ciliate; bracts
similar to bracteoles, in 2--3 pairs, 2-fid for 1/6--1/3 of the length or only
emarginated, dentate-ciliate in apical part. Androecia on short postical (more rarely lateral) branches, commonly
continuing vegetative growth, bracts in 4--5 pairs, ovate to subquadrate, bracteoles
2-fid, with two subulate lobes. Capsule
oval, ca. 900 x 500 \um; seta 10--15 mm long and 135--210 \um in diameter. Elaters ca. 10--12 \um thick, 2-spiral,
spirals reddish brown. Spores 10--15
\um, reddish brown, delicately papillose. Acidophilic to moderate
neutro-tolerant meso- hygrophyte
of partly to strongly shaded places, northward on moderately moistened peaty banks
of streams, southward on sandy-loamy banks of ditches, occasionally acid
rock; elevation 0--2200 m; B.C., N.S.; Ala., Alaska, Ark., Calif., Conn., Del.,
D.C., Fla., Ga., Ky., Maine, Mass., Mo., N.H., N.J., N.Y., N.C., Oreg., Pa., R.I.,
S.C., Tenn., Va., Wash., W.Va.; Eurasia. In the North Kurzia makinoana is
associated with open swampy massifs also near sea coast, although southward the
species seems to be frequent in the areas occupied by woody swamps (also evergreen).
Aside ‘swampy’ area this species growing over acidic rocks and over Sphagnum and other bryophytes in higher
elevations in the Appalachians, where also occurs in low elevations in forests
where confined to peaty and sandy banks of streams. 2. Kurzia pauciflora (Dickson) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 171, 1963
[1964] Jungermannia pauciflora Dickson, Fasc. Pl. Crypt. Brit. 2: 15, 1790; Kurzia setacea Grolle;
Microlepidozia setacea (Mitten)
Jørgensen Plants dull to deep green
and brownish green, rarely dark brown, in dense mats, 3--20(30) mm long, 200--300
\um wide, creeping to ascending and suberect. Stem 50--80 \um in diameter, more or less regularly pinnately or 2-pinnately
branched, with branches occasionally terminating in flagellae (become microphyllous
and rhizogenous). Rhizoids a few to
numerous, in ventral and lateral flagellae (microphyllous branches) common. Leaves transverse to incubous oblique,
mostly distant, subquadrate to obtrapezoidal when flattened,
180--250 x 180--250 \um, merely symmetrical, deeply 3--4(--6)-lobed, with undivided
area of 2--3 cells high, lobes entire, subulate to narrowly lanceolate, (2--)4--5 cells wide in base, dorsal lobe slightly smaller than
others; branch leaves commonly smaller than stem leaves and only 2--3-lobed. Underleaves similar, but smaller than
leaves, 3--4-lobed to 1--2 cells to the base, slightly convex, 115--225 \um long
and 65--120 \um wide. Leaf lobe cells
18--25 x 13--19 \um at lobe bases, with walls slightly thickened, cuticle slightly
verruculose. Perianth terminal, on
short postical branches, rarely also at apex of lateral branches, ovoid-cylindrical
to fusiform, basically 3-plicate, obtusely 4--7-plicate near mouth, mouth lobulated
to ciliate; bracts similar to bracteoles, in 2--5 pairs, 3- or 4-fid for 1/3--1/2
of the length, dentate-ciliate in apical part. Androecia on short postical branches, commonly continuing vegetative
growth, bracts in 2--5 pairs, ovate to subquadrate, 2--4-lobed, bracteoles 2-fid,
with two subulate lobes. Capsule ellipsoidal,
ca. 900 x 500 \um; seta 10--15 mm long and 135--210 \um in diameter. Elaters ca. 6--8 \um thick, 2-spiral,
spirals reddish brown. Spores 8--14
\um, yellowish brown, delicately papillose. Acidophilic to moderate
neutro-tolerant meso-hygrophyte
of partly to strongly shaded or rarely open places; elevation 0--700 m; B.C.,
Nfld. and Labr. (Nfld.), N.S., Nunavut; Alaska, Conn., Maine, Mass., Mich., N.H.,
N.J., N.Y., Ohio, Vt.; Eurasia. Kurzia pauciflora occurs as single shoots between Sphagnum and Leucobryum in mires, sometimes with Mylia anomala, Cladopodiella fluitans, Cephaloziella spinigera, and other liverworts.
Occasionally it is found on bare peat in heaths, wetlands and wet cliffs and stones
with Bazzania denudate or Herbertus aduncus. Unlike Kurzia makinoana,
this is species confined to Sphagnum
carpets in bogs with very acid reaction (below 3.8 pH).
It also avoids watercourse banks that K.
makinoana prefers. 3. Kurzia trichoclados (Müller Frib.) Grolle, Rev. Bryol. Lichénol.
32: 171. 1963[1964] Lepidozia trichoclados Müller Frib. , Hedwigia 38 (4): 197, 1899; Microlepidozia trichoclados
(Müller Frib.) Jørgensen Plants yellowish to pale
green and yellow brown, rarely dark yellowish green, in large dense patches, 20--60
mm long, 150--300 \um wide, creeping. Stem
80--110 \um in diameter, irregularly to regularly 2-pinnately branched, with branches
occasionally terminating in flagellae. Rhizoids
few to frequent in leafy stems, abundant in flagellae. Leaves transverse, distant to contiguous and subimbricate,
subquadrate when flattened, convex, 150--220 x 180--300 \um, deeply (for 0.6--0.8
of the length) 3--4-lobed, with undivided area of 2--3(--4) cells high, lobes
entire, subulate to narrowly lanceolate, 2--4 cells wide in base, dorsal lobe
slightly smaller than other; branch leaves commonly smaller that stem leaves and
only 2--3-lobed. Underleaves smaller
than leaves, (2--)3--4-lobed to 2--3 cells to the base,
slightly convex, 90--140 \um long and 140--190 \um wide, or sometimes as large
as leaves. Leaf lobe cells 20--30 x
14--18 \um at lobe bases, with walls slightly thickened, cuticle slightly striolate
verruculose. Perianth terminal, on
short postical branches, ovoid-cylindrical to nearly fusiform, basically 3-plicate,
mouth crenulate to ciliate; bracts similar to bracteoles, in 2--3 pairs, ovate
to oblong ovate, 2(--3)-lobed for 1/5--1/4 of the length, entire to denticulate
and shortly laciniate. Androecia on
short postical branches, commonly continuing vegetative growth, bracts in 4--20
pairs, ovate to subquadrate, bracteoles 2-fid, with two subulate lobes. Capsule ellipsoidal. Elaters 10--11 \um thick, 2-spiral. Spores 12--14.5
\um, yellowish brown, verruculose. Acidophilic to neutral-tolerant,
meso- to hygrophyte of partly shaded to open wet cliffs,
sometimes over and among Sphagnum in
cliff ledges and wetlands, rarely on decaying logs; elevation unknown, probably
below 500 m; B.C.; Eurasia 3. LEPIDOZIA (Dumortier)
Dumortier, Recueil Observ. Jungerm., 19. 1835, nom. conserv. * [Greek lepis. scale, and
ozos, branch, for the imbricate female bracts] Pleuroschisma sect. Lepidozia Dumortier, Syll. Jungerm. Europ.: 69. 1831 Plants prostrate to ascending,
in dense to loose patches, deep green to greenish never brown or red, strongly
varying in size (0.4--5 mm). Stem freely
pinnately to 2-pinnately (Frullania-type or postical-intercalary)
branched for normal branches, occasionally with rhizogenous flagellae; hyalodermis
absent, epidermal cells similar in size or slightly larger than inner. Rhizoids virtually absent or common, originated
near ventral underleaf base. Leaves
not deciduous, incubous, oblique, 3--5-lobed, symmetrical to slightly secund,
rather small, lobed for 1/3--4/5 of the length, lobes 3--6 or more cells wide,
triangular, margin entire to spinose-dentate. Underleaves similar in size to leaves or smaller, symmetrical, obliquely
to erect-spreading. Leaf lobe cells
mostly with thickened cell walls or walls more or less thin, trigones small to
vestigial; cuticle smooth to (rarely) papillose; oil bodies mostly smooth surfaced,
5--16 per cell. Sexual condition dioicous or monoicous, but commonly sterile.
Perianth on short postical branches,
3-plicate, elongate, long-conical to nearly fusiform, bracts in several pairs,
with distalmost circle of bracts larger than leaves, subentire to shallowly lobed.
Androecia as short postical branches
or terminal on short to elongated lateral branches that sometimes become flagellate;
male bracts similar to leaves, but more shallowly lobed, antheridial stalk biseriate.
Capsule oblong cylindric, capsule wall
3--5-stratose; with robust seta. Elaters
2-spiral. Spores brown, papillose. Species ca. 60 (3
in the flora): worldwide except Antarctic. SELECTED REFERENCES Hong,W. S. 1988. The family Lepidoziaceae
in North America west of the hundredth meridian. Bryologist 91:326--333. Schuster R. M. 1969. The Hepaticae and Anthocerotae
of North America, Vol. 2. New York, Columbia University Press. 1. Leaves narrower
than the stem, stem ca. 2/3 of the width of main axis
with leaves, leaves very distant … 3. Lepidozia sandvicensis 1. Leaves distinctly
wider than stem, stem ca. 1/2 or less of the width of
the main axis with leaves. 2. The largest leaf
lobe of the leaf 9--15 cells wide, leaves distinctly convex, dioicous … 1. Lepidozia filamentosa 2. The largest leaf
lobe of the leaf 4--8 cells wide, leaves nearly plane or slightly conves or curved to ventral side of the stem, autoicous …
2. Lepidozia reptans 1. Lepidozia filamentosa
(Lehman & Lindenberg)
Lehman & Lindenberg, Syn. Hepat.
2: 206. 1845 Jungermannia filamentosa Lehman & Lindenberg, Nov. Stirp.
Pug. 6: 29. 1834 Plants light green to green
rigid, loosely to distinctly ascending, 20--50 mm long, 800--1500 \um wide, more
or less rigid, forming loose pure patches or scattered among mosses. Stem 500--650 \um wide and 400--450 \um
high, easily pinnately to 2-pinnately branched, branches commonly transform to
flagelliform apex. Rhizoids virtually
absent in distal part of the shoots, but
merely common in older sectors and abundant in flagellae, originating near underleaf
bases. Leaves contiguous to subimbricate or more rarely distant, incubous, oblique, distinctly
to slightly convex, subquadrate to trapezoidal, nearly symmetrical to merely asymmetrical
distant, 700--900 \um long and 700--900 \um wide, 3--4-lobed for 1/3--1/2 of the
length, lobes triangular, slightly acute, ended by 1--2 superposed cells, or blunt,
7--15 cells broad in the base. Underleaves
ca. 2/3 of lateral leaves in size, subquadrate to trapezoidal, 3--4-lobed to 1/3--1/2
of the length, 350--450 x 650--750 \um. Leaf
lobe cells in their bases, subquadrate to hexagonal 20--50 x 20--30 \um, subequally thick-walled, or walls almost thin, trigones small,
concave, cuticle smooth; oil bodies 7--15 per cell, smooth-surfaced, elliptical
to fusiform, 4--5 x 2--3 \um. Sexual condition dioicous. Perianth fusiform, 4--6 mm long, 3-plicate
in distal half. Androecia on short
postical branches, with 2--5 pairs of bracts, monoandrous.
Capsule ca. 2 mm long, seta ca. 1 cm
long. Elaters 2-spiral, spores brown, papillose. Acidophilic mesophyte preferring decaying wood in oroboreal
to oro-hemiboreal dark coniferous forests; elevation
0--1000 m; B.C.; Alaska, Wash.; South America; Eurasia; Oceania. 2. Lepidozia reptans (Linnaeus) Dumortier, Recueil Observ. Jungerm.,
19. 1835 Jungermannia reptans Linnaeus, Sp. Pl. 1: 1133. 1753 Plants light green to green
prostrate to very loosely ascending, 15--40 mm long, 450--1000 \um wide, more
or less rigid, forming loose pure patches or scattered among mosses. Stem 200--300(--350) \um wide and 150--200
\um high, once pinnate or rarely 2-pinnate, branches sometimes with flagelliferous
apex. Rhizoids virtually absent in
distal part of the shoots, but merely common in older sectors and abundant in
flagellae, originating near underleaf bases. Leaves contiguous to subimbricate or rarely
distant, incubous, oblique, slightly convex of nearly plane, subquadrate to trapezoidal,
nearly symmetrical to merely asymmetrical distally, 500--650 \um long and 450--800
\um wide, 3--4-lobed for 1/3--1/2 of the length, lobes triangular, slightly acute,
ended by 1--2 superposed cells, or blunt and decurved,
4--8 cells broad in the base. Underleaves
ca. 2/3 of lateral leaves in size, quadrate to trapezoidal, 3--4-lobed to 1/3--1/2
of the length, 300--400 x 350--450 \um. Leaf
lobe cells in their bases, subquadrate to hexagonal 20--30 \um in diameter,
equally thick-walled, or walls almost thin, trigones vestigial to small, concave,
cuticle smooth; oil bodies 8--25 per cell, smooth-surfaced, spherical to oblong
and shortly fusiform, occasionally segmented to nearly botryoidal. Sexual condition
autoicous. Perianth long and prominent,
cylindrical-fusiform, 3-plicate distally, on short postical branches, bracts much
larger than leaves, similar to bracteole, broadly ovate. Androecia on short postical branches, bracts small and pellucid, in
4--8 imbricate pairs, strongly concave, monoandrous.
Capsule yellow-brown, (3--)4-stratose, seta with 12 epidermal and 16 inner rows of cells.
Elaters 9--12 \um in width, slightly
tapering at the ends, 2-spiral. Spores
11--15 \um, reddish to yellowish brown, verruculose. Acidophilic to neutral-tolerant
mesophyte, occurring in partly to full shaded sites
over various substrata from decaying wood in forests and humus along streams,
to cliffs and peaty sides of tussocks in tundra belt; elevation 0--2500 m; Alta.,
B.C., N.B., Nfld. and Labr. (Nfld.), N.W.T., N.S., Nunavut, Ont., P.E.I., Que.,
Sask., Yukon; Alaska, Calif., Colo., Conn.,
D.C., Idaho, Iowa, Ky., Maine, Mass., Mich., Minn., Mont., N.H., N.J., N.Mex.,
N.Y., N.C., Ohio, Oreg., Pa., R.I., Tenn., Vt., Va., Wash., W.Va., Wis., Wyo.;
Eurasia. Lepidozia reptans occurs on decaying wood associated with common xylophytes, like Scapania
apiculata, Riccardia palmata, Blepharostoma trichophyllum,
Anastrophyllum michauxii, Nowellia, etc. In
stones may grow with Diplophyllum taxifolium and
even Marsupella emarginata and
Tritomaria
quinquedentata,
although commonly forms pure patches. When growing on the sides of tussocks in
mires or on tundra, it is associated with Calypogeia muelleriana, Cephalozia bicuspidata, etc. By general distribution on North America,
it is a boreal species, rarely occurring in true tundra or in deciduous forest
zone. 5. Lepidozia sandvicensis
Lindenberg, Syn. Hepat. 2: 201, 1845 Plants pale green, in loose
mats, 20--50 mm long, 500--800 \um wide. Stem
400--500 \um wide and 200--300 \um high, easily pinnately to 2-pinnately branched,
branches commonly transform to flagelliform apex. Rhizoids virtually absent in distal part of shoots, but occasional
in flagellae tips. Leaves strongly
distant, subtransverse, distinctly to slightly convex, subquadrate, nearly symmetrical
to slightly asymmetrical, 200--300 \um long and ca. 400 \um wide, (2--)3--4-lobed for 1/2--2/3 of the length, lobes triangular, slightly
acute, ended by 1--2 superposed cells, or blunt, 2--6 cells broad in the base.
Underleaves ca. 2/3 of lateral leaves
in size, subquadrate to trapezoidal, 3--4-lobed to 1/3--1/2 of the length, ca.
250 \um wide. Leaf lobe cells in their
bases, subquadrate to hexagonal 22--35 \um in diameter, subequally
thick-walled, trigones small, concave, cuticle smooth. Sexual condition
dioicous. Perianth unknown. Androecia on short postical branches,
spicate, with 4--8 pairs of bracts, uni- to biandrous.
Capsule unknown. Elaters and spores unknown. Moist soil or moss
patches in moist coniferous forests; elevation 0--500 m; B.C.; Alaska; Central
America(?); Oceania (Hawaii). 4. TELARANEA Spruce
ex Schiffner, Hepat. (Engl.-Prantl), 103, 1893, nom.
conserv. * [Latin tela, plant tissue, and aranea, arachnoid,
alluding to thick and delicate branches, and leaves divided into filiform segments] Plants prostrate to ascending
or creeping, always delicate, among other bryophytes, pale to whitish green, pellucid,
lax and soft textured. Stem with distinct
hyalodermis formed by large thin- to evenly thick-walled cells, distinctly different
from medullary cells, loosely to densely pinnately to 2-pinnately branched, vegetative
branches terminal-lateral, sexual branches, at least in the most part, postical.
Rhizoids restricted to underleaf bases. Leaves
incubous to almost transverse, deeply and symmetrically 2--6(--13)-lobed, divided
almost to the base into subulate and filiform lobes (disc vestigial or very low).
Underleaves obliquely to erect-spreading,
similar to leaves and divided into 2--8 uniseriate cilia. Lobe cells elongate (2--4.5 as long as
wide), leptodermous, collapsed when dry, with smooth
cuticle. Sexual condition dioicous or autoicous, with archegonia and androecia
appearing on short ventral intercalary branches. Perianth obtusely 3-plicate, setose at the
mouth, developed on short postical branches, perigynium absent; female bracts
similar to bracteoles, varying from laciniate to shallowly 3--4-lobed, imbricate.
Androecia with antheridia solitary,
stalk 1-seriate. Capsule ovoid, capsule
wall (2--)3--5-stratose; seta of 8 epidermal and numerous
internal rows. Elaters 2-spiral, rarely
3-spiral, with blunt apices. Spores
small, 1--1.5 of elater diameter, finely, but distinctly areolate. Species ca. 100 (1
in the flora): North America, Mexico, Central America, South America,
Eurasia, Africa, Oceania. SELECTED REFERENCE Schuster R.
M. 1969. The Hepaticae and Anthocerotae of North America, Vol. 2. New York, Columbia
University Press. 1. Telaranea nematodes (Gottsche ex Austin) M.A. Howe, Bull.
Torrey Bot. Club 29(5): 284. 1902 Cephalozia nematodes Gottsche ex Austin, Bull. Torrey
Bot. Club 6 (52): 302. 1879 Plants pale to whitish green,
when dry mostly collapsed, prostrate, very delicate, 5--10 mm long, 300--450 \um
wide, among and over other bryophytes. Stem
120--150 \um wide and 90--105 \um high, with 10--12 hyalodermis cells rows, cortical
cells 40--45 \um wide, walls somewhat thickened both inside and in outer layer,
easily branched. Rhizoids a few, in
erect to obliquely spreading short fascicles originating near underleaf bases.
Leaves distant to contiguous, divided
to nearly base, disc 1(--2) cells high, with 3--4 uniseriate (robust biseriate
at the bases) lobes, obliquely spreading, with cilia 4--8 cells long, 350--600(--900)
\mu long. Underleaves smaller than
leaves, 2--3-lobed nearly to the base, incurved distally. Cilia cells 90--110 x 20--28 \um, thin-walled,
trigones vestigial, cuticle smooth to faintly papillose; oil bodies homogenous,
nearly spherical, minute, several per cell. Sexual condition autoicous.
Perianth terminal, on short postical
branches, rarely on elongate lateral branches, long ciliate, with cilia commonly
2--3-furcate at the mouth, bracts in 2--3 pairs, with long tapered cilia or laciniate.
Androecia on short lateral branches,
bracts similar to vegetative leaves, monandrous. Capsule wall 3-stratose, epidermal cells
narrowly rectangular, with nodular thickenings, inner cells narrowly rectangular
with several bands of thickenings. Elaters
ca. 180--240 \um long and 9--13 \um thick, 2(--3)-spiral, spirals red-brown. Spores 14--17 \um, pale brown, with areoles
1--1.8 \um in diameter. Acidophilic meso-hygrophyte, frequent in coastal swamps associated with
Magnolia, Persea, Gordonia, etc., where growing along
sluggishly flowing streams on bare sandy ground and creeping over Sphagnum; elevation unknown, probably below
1000 m; Ark., Fla., Ga., Miss.,
N.J., N.Y., N.C., S.C., Tex., Va.; Mexico; Central and South America;
Eurasia; Africa; Oceania. Telaranea nematodes is commonly associated with the liverworts like Pallavicinia, Odontoschisma, Cephalozia, more rarely with Microlepidozia, Calypogeia and other
hygrophytic liverworts. Common in coastal swamps although
sometimes occurred as far as over 300 miles inward of the continent (in North
America). |
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