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BFNA
Title: Lophocoleaceae |
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XX.
LOPHOCOLEACEAE Vanden Berghen John J. Engel Plants forming mats. Branches mostly of
lateral-intercalary type; Frullania type branches less common; Andrewsianthus
type (from dorsal angle of leaf) present in Evansianthus;
ventral-intercalary branches much less frequent, present in some taxa. Rhizoids,
typically from stem in a sharply defined region at underleaf base, in Leptophyllopsis
from cells of underleaf. Leaves alternate to subopposite, with
insertion typically strongly succubous to at times nearly horizontal, unlobed
or often 2-lobed, the margins often entire or subentire, occasionally with
scattered accessory teeth or cilia. Underleaves typically large and
conspicuous, inconspicuous in a few taxa, usually free or narrowly connate on
one side, exceptionally distinctly connate on both sides, typically 2-fid to
varying degrees, at times emarginate, the lamina margins mostly each bearing
1--several teeth or laciniae, at times entire. Specialized
asexual reproduction occasional, by regeneration in situ from adult
leaves, by gemmae individually produced from leaf margins (never catenate),
by caducous leaf segments, or by caducous leaves Gynoecium
triquetrous or not, with position similar to that of androecia; bracts often
larger than leaves, often +/- 2-lobed, less often truncate-emarginate, the
margins entire or at times with accessory armature; bracteole typically large
and similar to bracts, exceptionally small and inconspicuous, usually free or
narrowly connate on one side; vestigial stem perigynium or coelocaule
precursor sometimes present. Perianth long-exserted, symmetrically
trigonous or with the ventral face narrower and at times also shorter, the
ventral face narrow and often folded inward, with the mouth appearing
bilabiate, the perianth with 2 keels lateral, the third dorsal (the keels at
times with a wing), occasionally inflated to the point where keels are
obscure; mouth 3-lobate, wide and open, the lobes undivided or 2-lobed,
entire or often spinose-dentate or ciliate.
Genera 24 (3 in the flora),
species ca. 400 (12 in the flora). The family Lophocoleaceae is
predominantly south-temperate-subantarctic and tropical. In non-tropical
parts of the Northern Hemisphere representation is sparse, and the relatively
few taxa all belong to the either Chiloscyphus, Cryptolophocolea or Leptoscyphus. Traits of
the family supplemental to the description above follow: Plants usually
moderately to distinctly anisophyllous, subisophyllous to isophyllous in a
few taxa, usually prostrate, occasionally suberect to erect, whitish to clear
to yellow-green or developing brownish to chestnut-brown to fuscous
pigmentation, small to medium to large. Stolons are present in some taxa. Stems
with cortical cells not strongly differentiated, but well defined and in 1--3
rows of thick-walled cells in a few taxa, a hyaloderm is absent. Rhizoids usually colorless, rarely
brownish, usually common. Leaves mostly uni-stratose, polystratose in a few
taxa. Cells thin-walled, with trigones minute to large to strongly nodular;
surface usually smooth, infrequently striate-papillose or tuberculate with
1 protuberance per cell, hispid in a few species. Oil-bodies conspicuous, usually 2--10 per
cell, usually papillose to weakly to distinctly botryoidal. Sexual condition dioicous, autoicous, or
paroicous. Androecia on main shoot
or rather short to moderately long Frullania type, lateral-, or
ventral-intercalary branches (and then often terminal in position but
becoming intercalary), or on determinate, compactly spicate
lateral-intercalary branches lacking normal leaves; bracts at least
moderately differentiated from leaves, saccate at least at dorsal base to
form a +/- defined lobule, the free margin of lobule often with an incurved
tooth; antheridia mostly 1 per bract, the head subglobose, with cells
irregularly arranged, the stalk 1--2(rarely 4)-seriate. Seta typically with numerous cell rows. Capsule oblong-ovoid, exceptionally
long-elliptic to cylindrical, the valves straight, not twisted, (3)4--6(8)-stratose,
the outer layer of cells typically equal in thickness to ca. 1.5 of interior
strata; outer layer of cells with relatively small to large nodular
thickenings on radial walls (the outer layer of cells devoid of thickenings
and hyaline in Heteroscyphus spp.); innermost cell layer of cells with
semiannular bands, the bands at times incomplete, exceptionally forked and
anastomosing to delimit well-defined fenestrae (Cryptolophocolea
spp.). Spores papillose grading to
short-vermiform ridges, the ridges at times uniting to form areolae,
occasionally echinate; spore to elater diam. ratio averaging 1.1--2.8:1. Elaters free, usually 2-spiral. SELECTED REFERENCES: Schuster, R. M. 1980. The Hepaticae and Anthocerotae of North
America east of the hundredth meridian.
Vol. IV. Columbia University
Press, New York. 1. Plants brown to fuscous; perianth
bilabiate, laterally compressed at least above
................................... 1.
Leptoscyphus, p. XXX 1. Plants whitish to pure green; perianth
trigonous, not laterally compressed. 2.
Leaves alternate (rarely
subopposite), free dorsally, never connate, the apex undivided or
fundamentally 2-fid, the lobes then +/- subequal; underleaves free or
narrowly connate on one or (occasionally) both sides; antheridial stalk
uniseriate (rarely multiseriate); capsule wall inner layer with semiannular
bands, if (occasionally) forking and anastomosing then with fenestrae few and
sporadic or altogether lacking (except C. perpusillus)
................................................... 2. Chiloscyphus, p. XXX 2.
Leaves subopposite to opposite,
free or connate dorsally, the apex short-2-fid, the ventral lobe mostly
larger than the dorsal; underleaves typically broadly connate on both sides;
antheridial stalk biseriate; capsule wall inner layer with semiannular bands
often forking and anastomosing to delimit fenestrae ........... 3. Cryptolophocolea, p.
XXX 1.
LEPTOSCYPHUS Mitten, J. Bot. Kew Garden Misc. 3: 358. 1851 * [Greek leptos, thin, slender, and skyphos,
cup, alluding to the flattened perianth] Laura Briscoe John J. Engel Anomylia R. M. Schuster Plants pure green to
olive-green, with exposure usually becoming golden brown to deep brown to
reddish brown, rarely piceous or fuscous, medium to vigorous in size, rarely
small. Stems with cortex poorly to moderately differentiated, with
cortical cells in 1--2 layers of somewhat smaller or somewhat larger cells,
the cortical cells with walls often thicker than those of the medullary;
branching irregular, lateral-intercalary, or less commonly Frullania
type or ventral-intercalary type. Rhizoids hyaline, exceptionally rose
to reddish brown, in fascicles from stem at base of underleaves, at times
becoming scattered along ventral surface of stem, particularly in older shoot
sectors. Leaves with orientation
slightly [to strongly] succubous, the lines of insertion extending to stem
midline dorsally or, in some species not extending to stem midline and then
delimiting a leaf-free strip of stem cells; leaves alternate to opposite,
usually free dorsally, exceptionally connate dorsally, convex to concave,
undivided and entire or 2-lobed [the apex rarely with 3--4 long cilia];
dorsal margin entire [or rarely with several teeth or cilia]; ventral margin
entire [or with a tooth or lacinia in distal half or 1--3 dentate-laciniate
near base or ciliate]. Cells
thin-to somewhat thick-walled, with trigones well defined, medium and
straight-sided to nodose; surface smooth or wax-like, the surface strongly
papillose in a few species, rarely striolate toward leaf base. Oil-bodies 2--7 up to 14 per median
leaf cell, hyaline or grey, granular to papillose to coarsely
botryoidal. Underleaves (0.5--)1--3 times or more stem width, connate on both
sides, occasionally connate only on one side, rarely free, mostly 2-fid to
0.5--0.9, or, occasionally, shallowly 2-lobed or 2-dentate or undivided and
lanceolate, the lobes acuminate to caudate; lamina margins on each side
usually with a spreading tooth or cilium or lacinium, at times with accessory
small teeth. Specialized asexual
reproduction via caducous leaves lacking. Sexual condition dioicous or monoicous. Androecia terminal but becoming intercalary, subfoliose to +/-
spicate; bracts similar to leaves except smaller, ventricose in basal
portion, the lobule margin involute, entire or with a few small teeth;
antheridia 1(rarely 2) per bract, the stalk 2(--4)-seriate, exceptionally
uniseriate. Gynoecia terminal on
leading axes or on short intercalary branches; bracts slightly to
considerably larger than leaves, usually similar to leaves in shape,
undivided or 2(--4)-lobed, the margins entire or with a few small teeth, in a
few species with cilia; bracteole (0.01--)0.2--0.6 times area of bracts,
typically much larger and usually longer than underleaves, connate on 1 or
both sides or free, 2--4-lobed, rarely undivided (L. cuneifolius), the
margins entire or sparingly dentate to laciniate, with a few papillae,
exceptionally ciliate. Perianth rather inflated basally,
bilaterally compressed at least distally, the lateral lobes prominent, the
ventral lobe narrower and often shorter, often infolded and narrowly sulcate,
the perianth thus feebly, unequally 3-lobate, the mouth wide and truncate,
entire or dentate to spinose to ciliate; keels narrowly rounded, at times
rather sharp, with wings often present on one or more keels or between the
keels. Seta of the general type,
comprised of many rows of +/- similar cells or (L. cuneifolius) with 8--10 outer + 4 rows of inner cell. Capsule
wall 2(--3) stratose [to 4--7(--8)-stratose]; outer layer of cells typically
with well-developed nodular thickenings; innermost layer of cells with
semiannular bands well developed or (in a few species) lacking and merely
with nodular thickenings. Elaters
2-spiral. Spores various:
vermiculate or papillose-vermiculate or bacculate or echinate. Species ca. 33 (1 in the
flora): North America, West Indies,
Central America, South America, Europe, Asia, Africa, Atlantic Islands
(including Macaronesia), Pacific Islands (including Hawaii and New Zealand),
Australia. The
bilabiate, laterally compressed perianths and the brown to fuscous
pigmentation will distinguish Leptoscyphus
from other genera of the family. The
genus was monographed by R. Grolle (1962).
Leioscyphus Mitten in Hooker
is an illegitimate synonym. SELECTED
REFERENCES: Grolle, R. 1962. Monographie der Lebermoosgattung Leptoscyphus Mitt., Nova Acta Leop.
25: 1---141. Vanderpoorten, A., A.
Schäfer-Verwimp and J. Heinrichs. 2010.
The taxonomy of the leafy liverwort genus Leptoscyphus (Lophocoleaceae) revisited. Taxon 59:
176---186. 1. Leptoscyphus cuneifolius (Hooker) Mitt., Hooker’s J. Bot.
Kew Gard. Misc. 3: 358. 1851 Jungermannia cuneifolia Hooker, Brit. Jungerm., 64.
1814; Mylius cuneifolius (Hooker)
S. Gray Plants filiform, brown to reddish brown
to blackish brown, the shoot tips green or yellowish green, the shoots
minute, ca. 300--450(--650) /um wide, 2--8 mm long. Stems
comprised of relatively few, thick-walled cells, the cortical cells slightly
larger and thicker-walled than those of the medulla; branches of Frullania type;
ventral-intercalary branches also present.
Rhizoids short, sparse,
fasciculate, from stem at base of underleaves. Leaves
slightly obliquely succubous, the insertion narrow, short, +/- straight, the
lines of insertion not extending to stem midline dorsally and delimiting a
leaf-free strip of 2 cells wide; leaves fragile and freely caducous, remote,
widely spreading to squarrose, often slightly concave (dorsal view), broadly
obovate to broadly cuneate, the width at the subapical sector equal to the
length or slightly wider, the apex broadly rounded-truncate to
truncate-retuse, the margins entire. Cells of leaf with walls thin to
somewhat thickened, with trigones large to bulging, the median cells
subisodiametric, (12--)16--18(--20) /um wide x 18--22(--25) /um long; surface
smooth. Oil-bodies (Schuster, 1980) 2--3, occasionally 4--5 per cell,
colorless, ovoid to short-ellipsoid or spherical, ca. 4.5--6 to 4 x 5--6.5
/um up to 6 x 9 /um. Underleaves polymorphic, ca. as wide
as stem, much smaller than leaves, free or narrowly connate on 1 side,
undivided, variable in shape, lanceolate to ovate-triangular or cuneate to
subulate, rarely 2-lobed. Specialized asexual reproduction by
caducous leaves, the shoots at times entirely devoid of leaves. [Sexual
condition dioicous. Androecia terminal but becoming
intercalary in position; bracts in up to 4 pairs, imbricate, strongly
ventricose; lobule large for bract size, the lobule strongly tapering
distally and forming an acute to lanceolate process; antheridia solitary, the
stalk biseriate. Gynoecia terminal
on leading shoots; bracts much larger than leaves, subrotund, undivided and
entire; bracteoles ca. 0.5 times bract size, much larger than the
underleaves, free, oblong ovate, undivided, the apex obtuse, entire, the
margins on each sides occasionally with obtuse teeth. Perianth
oblong, the base inflated, conspicuously laterally compressed distally; mouth
with large, irregular, subacute to blunt teeth. Seta 4 cells, with 8--9(--10) rows of outer cells surrounding an
internal core of 4 cells. Capsule
short ovoid, the wall 2--3-stratose, the outer layer of cells with nodular
thickenings on longitudinal and transverse walls, the nodules often dilated
distally; innermost layer of cells with short, narrow, parallel-sided
thickenings, with semiannular bands completely absent. Spores echinulate-hispiduous.] Usually
epiphytic, also moist rock, high precipitation areas of Southern
Appalachians; high elevations; N.C., Tenn, Va.; West Indies (Dominican
Republic, Dominica, Guadeloupe, Jamaica, Martinique); Central America (Costa
Rica, Panama); South America (including Galapagos Islands, Juan Fernandez);
Europe (Ireland, U.K., Portugal, Norway); Eurasia (Turkey); Atlantic Islands
(Tristan da Cunha, Macaronesia). Leptoscyphus cuneifolius rarely forms pure colonies. In
the flora it is usually associated with other hepatics including Bazzania nudicaulis, Plagiochila exigua, and Sphenolobopsis pearsonii. Regionally,
this species is limited to high elevation mountain summits dominated by red
spruce-Fraser fir forests. It occurs
on the bark of Abies fraseri and is
apparently restricted to asexual reproduction 2, CHILOSCYPHUS Corda in Opiz (ed.), Beiträge zur
Naturg. (12): 651. 1829 as Cheilocyphos; corr. Dumortier, Sylloge
Jung. Europ. Indig. 67. 1831, nom. et orth. cons. * [cheilos, lip, edge, and kyphos,
bent, alluding presumably to the shape of the mouth of the perianth] John J.
Engel Lophocolea (Dumortier) Dumortier, Recueil Observ. Jungerm.
17. 1835 Plants prostrate to erect, various shades of green and, less
frequently, brown, the main shoots of smaller species to 0.8 mm wide, of
larger species to 5 mm wide. Branching variable, the branches of
lateral-intercalary and Frullania
types, both common; ventral-intercalary branches occasional; stolons
absent. Stems poorly to rather well differentiated, with cortex in 1--2
rows or with cortex hardly differentiated.
Rhizoids colorless, from
stem at bases of underleaves, rarely becoming scattered, rarely septate. Leaves
alternate, rarely subopposite, free dorsally, the orientation mostly strongly
succubous, occasionally subtransverse, the insertion strongly succubous, not
or slightly to distinctly recurved at ventral end, the lines of insertion
mostly extending to stem midline dorsally, in some species not extending to
stem midline dorsally and then a leaf-free strip of stem cells present.
Leaves mostly convex, occasionally concave but never conchiform concave; apex
undivided or fundamentally 2-fid; margins entire or variously armed with
teeth, cilia or laciniae. Cells of leaf with walls thin,
trigones variable, minute to medium to distinctly knot-like; surface of
leaves smooth or, in some taxa, tuberculate or hispid. Oil-bodies
(1)2--7(--12) per cell, mostly coarsely papillose to coarsely botryoidal,
occasionally sharply granular. Underleaves 0.7--3.7X stem width,
free or narrowly connate on 1 or (occasionally) both sides, the insertion
crescent to inverted U-shaped, the underleaves plane to convex or concave,
with apices mostly 2(4--6)-lobed to at least 0.4, at times undivided and
entire or 2-dentate or retuse to 2-fid to 0.3, the lobes, when present,
commonly acuminate to subcaudate and with margins entire or variously armed
with teeth, cilia or laciniae; lamina margins entire or dentate to laciniate
to lobate. Asexual reproduction by caducous leaf lobes, regeneration or
gemmae that are not produced in fascicles.
Sexual condition dioicous,
autoicous or paroicous. Androecia on main shoot or, if on
short, abbreviated branches, then never consistently so; bracts similar in
size to vegetative leaves grading to much smaller, the dorsal base dilated
and forming an infolded flap or lobule; lobule margin commonly with a distal
process and, often, sparingly to densely armed with teeth and cilia;
antheridia solitary, rarely 2--3 per bract, the stalk usually uniseriate,
rarely multiseriate. Gynoecia on main shoot or elongated
branches or short, abbreviated lateral- or ventral-intercalary branches,
rarely on short, abbreviated Frullania
type branches; bracts of innermost series with apices undivided to retuse or 2-fid,
the margins entire or dentate to ciliate to laciniate; bracteoles of
innermost series conspicuous, mostly 0.5--1X bract area, free or connate on 1
or both sides, the apices undivided and entire or 2-dentate or retuse or 2--4-fid
to 0.1--0.7; lamina margins entire or dentate-laciniate-lobate. Calyptra
various, grading from ca. 0.5X perianth length to long emergent from perianth
apex. Perianth mostly sharply trigonous,
rarely terete, the mouth 3-lobed, the lobes +/- subequal or the ventral
slightly narrower, the lobes undivided and dentate to laciniate or entire or
shallowly to deeply 2-fid; keels often winged. Seta
with epidermal cells not greatly differentiated from the scattered, mostly
numerous inner cells. Capsule mostly ellipsoid, rarely long
ellipsoid to cylindrical, of (2--3)4--6 layers, the outer layer of cells
equal to thickness of at least 1.5--3.5 of interior strata or equal to or
exceeding the thickness of all interior strata combined, the outer layer of
cells with thickenings mostly on longitudinal radial walls and less frequent
or lacking on transverse walls, the thickenings nodule-like to spine-like;
innermost layer of cells with semiannular bands extending from radial walls,
the bands sometimes incomplete, sometimes forked but only occasionally
anastomosing and then not delimiting fenestrae or with fenestrae at most few
and sporadic, the radial walls often with nodule-like to spine-like
thickenings. Spores with dense, low, close papillae and short-vermiculate
markings. Elaters 2-spiral. Species
ca. 250 (10 in the flora, plus 2 subsp., 4 varieties): [[distribution]] Chiloscyphus is the largest genus in the family and has an
estimated 250 species. The genus, as recognized here, includes Lophocolea. Plants of this genus (including Lophocolea) are notorious for their
pronounced plasticity and difficulty.
The genus is well-represented in the south temperate. The treatment here is adapted and modified
from that of R. M. Schuster (1980). 1. Dorsal surface of leaf densely hispid;
underleaf segments with opposing teeth or cilia; perianths armed on external
(and often internal) surfaces with spinose cellular processes
...................…… 8. Chiloscyphus muricatus 1. Dorsal surface of leaf
uniformly smooth; underleaf segments entire, or at most with 1--2 teeth, the
teeth never regularly opposing; perianths smooth, never armed with spinous
processes (at least between keels). 2. Perianths restricted to
abbreviated lateral branches lacking normal leaves, the perianths short,
inflated; calyptra typically extending to
perianth apex or exserted slightly beyond it; androecium undifferentiated,
the bracts like normal leaves except slightly saccate at dorsal base. 3. Cells small: marginal cells of leaf apex (20--)26--28 /um,
where subquadrate; median cells ca. (21--)24--30 x 30--35(--40) /um (in
aquatic phases the cells often even smaller); oil-bodies fusiform to oval,
(1)2, more rarely 3--4(5) per cell, large:
4--5 x 7--12 /um to 6--7 x 8--12 (18) /um; plants green or dull to
deep green; lobes of perianth mouth not or hardly dentate; calpytra usually
long emergent from perianth apex ……………
9. Chiloscyphus
polyanthus 3. Cells larger: marginal cells of leaf apex (28)30--35 /um;
median cells (32)35--40 x 45--60 /um or even larger; oil-bodies mostly
(3--4)4--6(10--12) per cell, smaller:
4--5.5 /um and spherical to
4--4.5 x 8--9 µ,and ovoid,
rarely to 5 x 10--11 /um; plants usually whitish green; lobes of perianth
mouth usually +/- distinctly dentate; calpytra little or hardly emergent from
beyond perianth …………… 10. Chiloscyphus pallescens 2. Perianths typically terminal on main shoot
or leading branches, usually with at least some normal leaves, the perianths
typically elongate, not inflated; calyptra hidden deeply within the perianth,
not extending to perianth apex; androecium differentiated into a lobe and
ventricose lobule at dorsal base. 4. Mature leaves predominantly +/-
rectangular, often somewhat narrowed at base, the apices with lobes obtuse to
acute, or rounded, or some leaves entire (leaf lobes never terminating in a
uniseriate row of 3--7 cells; plants small, 0.5--1.6 mm wide; underleaves
uniformly free; paroicous; able to form gemmae (Chiloscyphus minor); plants +/- malodorous, yellow-green. 5. Abundantly fertile; gemmae
rarely developed (when present various, never of globular cell masses);
leaves at least of apices of mature stems entire or subentire; plants larger,
1--1.6 mm wide x 1--3 cm long
………………………. 6. Chiloscyphus profundus 5. Rarely with perianths; gemmae
freely developed, in the form of irregularly globose cell masses, rendering
the leaf margins erose; leaves (exc. where ragged because of gemmae
development) uniformly 2-dentate; plants small, 0 .5--0.8 mm wide x 10--18 mm
long …………………… 7. Chiloscyphus minor 4. Leaves +/- ovate or
ovate-triangular, widest near base and narrowed for most of their length, the
apices sharply 2-dentate or 2-lobed apices (the lobes rarely obscure; leaves
then subrotundate); leaf lobes terminating in a uniseriate row of 3--5(--7)
cells; plants larger, 1.5--3.5 mm wide usually; underleaves sometimes
narrowly connate on 1 side (rarely both sides) with leaf bases; dio- or
autoicous; unable to form gemmae; plants fragrant, whitish green to deep
green. 6. Leaf insertion lines not extending to stem
midline dorsally, delimiting a leaf-free strip of 1--2 cells wide; perianth
(known only for Chiloscyphus latifolius)
typically keels lacking wings. 7. Leaf marginal cells strongly crenate due to
conspicuously bulging cells, the lobe tips formed of moniliform cells; median
leaf cells often larger, 33--36(--40) /um wide; plants autoicous
.......................................
4. Chiloscyphus appalachicolus 7. Leaf marginal cells not bulging, the lobe
tips not of moniliform cells; median leaf cells smaller, 23--34 /um wide;
plants dioicous ......................................... 1. Chiloscyphus latifolius 6. Leaf insertion lines extending
to stem midline dorsally, a leaf-free strip lacking; perianth keels with
wings well developed, often high and armed with cilia and teeth. 8. Leaves polymorphous, varying from
suborbicular to ovate or feebly
oblate, the apex variable, undivided and entire to 1-dentate to 2-dentate to
weakly 2-lobed; female bracts with apex varying from 2-dentate to 2-dentulate
to 1-dentate to often subentire; male bracts rounded at the apex .................... 5. Chiloscyphus appalachiana 8. Leaves narrowly ovate to ovate-triangular,
clearly, sharply and uniformly 2-lobed, with narrowly acute to acuminate to
subcaudate lobes; female bracts distinctly 2-lobed, the lobes usually
narrowly attenuate; male bracts (unknown for C. coadunatus) 2-lobed or 2-dentate at the apex. 9. Underleaves typically free, sporadically
connate on 1 side; plants autoecious; perianth more shallowly lobed, the
lobes free for ca. 3--0.35 the perianth length .................. 3. Chiloscyphus cuspidatus 9. Underleaves connate on 1 or
both sides; plants dioicous; perianth more deeply lobed, the lobes free for
ca. 0.4--0.6 the perianth length .................................... 2. Chiloscyphus coadunatus 1.
Chiloscyphus latifolius
(Nees) J. J. Engel & R. M. Schuster, Nova Hedwigia 39: 418. 1985 (1984)
Lophocolea
latifolia Nees,
Naturg. Eur. Leberm. 2: 234. 1836; Jungermannia bidentata Linnaeus; Lophocolea bidentata (Linnaeus) Dumortier
Plants delicate, procumbent, aromatic when fresh, pale to whitish
green, translucent, 2.5--3 mm wide, 2--5 cm long. Branching
sparingly developed, the branches mostly of Frullania type, on average with less than 2 branches per cm of
stem; lateral-intercalary branches occasional. Rhizoids
sparingly developed. Leaves nearly horizontally oriented
and inserted, the lines of insertion
not extending to stem midline dorsally,
delimiting a leaf-free strip of 1--2 cells wide; leaves strongly
alternate, widely spreading, convex, asymmetrically rhomboid-ovate, widest
basally and distinctly narrowed toward the apex; apex narrow, nearly uniformly
2-lobed to 0.25--0.35, isolated leaves 3-lobed; lobes frequently divergent,
lanceolate-acuminate, terminating in a single cell or a uniseriate row of
2--5 weakly to moderately elongated, nonmoniliform cells; sinus usually
lunate; dorsal and ventral margins weakly to moderately arched, entire, not
crenulate, the dorsal margin long-decurrent.
Cells of leaf never tumid,
the walls thin, with trigones lacking, the median cells 23--34 /um wide,
30--35 /um long; surface smooth. Oil-bodies 2--5 per cell, in isolated
cells more numerous, finely granular-segmented, subspherical to
elongate-ovate, ca.. 4 x 4--8 /um. Underleaves about as wide as stem,
free or very narrowly connate on 1 side, the insertion shallowly inverted
U-shaped, the underleaves distant, plane or weakly concave toward base (ventral
view), narrowly rectangular to obdeltoid-subcuneate, 2-fid to ca. 0.5--0.8;
lobes acuminate to subulate-acuminate; lamina margins normally on each side
with a dentiform to acuminate tooth. Specialized asexual reproduction
absent. Sexual condition dioicous, usually sterile. Androecia
usually terminal, eventually becoming intercalary in position, spicate;
bracts in several pairs, smaller than leaves, asymmetrically 2--3 lobed. Gynoecia
on main shoot, elongate branches or short lateral-intercalary branches;
bracts larger than leaves, oblong-ovate, 2-lobed to 0.15--0.2 (fide Schuster
descr. ), the lobes acuminate, entire,
the dorsal margin of lamina weakly to moderately reflexed, not forming a
cnemis, entire, the ventral margin entire or with a tooth; bracteoles
narrowly connate on 1 side, ca.
0.5--0.8X length of bracts, concave,
2-lobed to ca. 0.25, the lobes entire, the lamina margins sharply reflexed,
entire or with a small tooth in distal sector. Perianth rarely developed, oblong, strongly trigonous, the mouth shallowly lobed, the lobes free for ca. 0.25--0.35 the perianth length, the
lobes conspicuously narrowed toward apex, the margins reflexed distally
lending the lobes +/- canaliculate toward the apex, each lobe shallowly 2-fid,
the segments narrowly acute to acuminate, coarsely dentate to laciniate, the
lobes otherwise coarsely dentate below level of segments; keels sharp, with
wings typically lacking, rarely present, then notably high, dentate. Capsule
ovoid to oblong-oval, the wall 3--5 layered; outer layer of cells with strong
nodular thickenings; inner walls with numerous semiannular bands. Spores
pale brown, 15--18 /um, nearly smooth.
Elaters 8--10 /um wide, 2-spiral.
Chiloscyphus latifolius differs from C. coadunatus and C. cuspidatus by the leaf insertion lines not extending to stem
midline dorsally and delimiting a leaf-free strip, and the typical lack of
perianth wings. For further comments
on C. latifolius, C. cuspidatus and
C. coadunatus see J. Vana and J. J.
Engel (2013); these concepts are followed here. On moist
soil, sometimes in silted mats indicating partial submersion; low to moderate
elevations; B.C.; Miss., N.Y., N.C., Ohio, Tenn., Va.; Europe. 2. Chiloscyphus coadunatus (Swartz) J. J. Engel & R. M. Schuster, Nova Hedwigia
39: 413. 1985 [1984] Jungermannia
coadunata Swartz,
Fl. Ind. Occid. 3: 1850. 1806 Plants soft and spongy, loosely
creeping, pale whitish green, to 2 mm wide (when leaves spreading), 2.1 cm
long. Branching rather common, the branches of Frullania- and
lateral-intercalary types. Rhizoids sparingly developed. Leaves strongly horizontally oriented and
inserted, the lines of insertion
extending to stem midline dorsally, a leaf-free strip lacking; leaves strongly alternate,
dorsally assurgent at least in basal half, often throughout, convex,
asymmetrically rhomboid-ovate, widest basally and distinctly narrowed toward
the apex; apex narrow, 2-lobed to 0.1--0.2; lobes parallel to weakly
divergent, narrowly acute to acuminate, terminating in a uniseriate row of
2--3 typically isodiametric cells (sporadically 1.5:1); sinus usually lunate;
dorsal and ventral margins weakly to
moderately arched, entire, not crenulate, the dorsal margin rather
short-decurrent. Cells of leaf never tumid, the walls thin, with trigones lacking,
the median cells 00—00 x
00--00 /um. Oil-bodies not seen. Underleaves
about as wide as stem, very narrowly connate on 1 or both sides, the
insertion shallowly inverted U-shaped, the underleaves distant, plane or
weakly concave toward base (ventral view), narrowly rectangular to obdeltoid,
2-fid nearly to base, the lamina 2--4 cells high between sinus base and
underleaf base; lobes acuminate to subulate-acuminate, entire, occasionally
with a small tooth; lamina margins normally on each side with a dentiform
tooth. Asexual Reproduction absent.
Sexual Condition
dioicous. Androecia not seen. Gynoecia on short ventral-intercalary
branch lacking normal leaves or on
main shoot; bracts somewhat Plagiochila-like in appearance, larger than leaves, erect and appressed to the perianth, strongly convex, oblong-ovate, elobate, the apex truncate,
with a few unicellular teeth, the dorsal margin of lamina strongly reflexed to revolute and forming a
distinct cnemis, entire, the ventral
margin entire; bracteoles narrowly
connate on 1 or both sides, +/- as long as bracts or somewhat shorter,
distinctly concave, obovate, 2-lobed to ca. 0.2--0.35, the lobes, entire, the
lamina margins sharply reflexed, entire or with a tooth. Perianths long linear, straight-sided, strongly trigonous, the mouth deeply lobed, the lobes free for ca. 0.4--0.6 the perianth length, the
lobes conspicuously narrowed toward
apex, the margins plane or weakly
reflexed distally, the lobes not
canaliculate toward the apex, each lobe
shallowly to more deeply 2-fid (to 0.5), the segments long attenuate,
sometimes acute, gradually tapering to 7--9 biseriate tiers and terminating
in a uniseriate row of 7--9 cells, the
uniseriate row sometimes with a few 1--3-celled teeth, the segment margins
otherwise entire, the lobes otherwise with to ca. 6 short to long cilia below
level of sinus base between segments, the cilia each with a uniseriate row
of 6--11 cells that are constricted at the septa; keels sharp, with wings well developed, notably high, up to 7 cells high, undulate, the wing margins with long cilia (type)
and several small teeth, rarely
entire. Sporophyte not seen. Habitat
unknown; moderate elevations; N.C.; West Indies (Cuba, Jamaica); South America
(Brazil), widely reported elsewhere in the Neotropics, but the reports
require confirmation. 3.
Chiloscyphus cuspidatus (Nees)
J. J. Engel & R. M. Schuster, Nova Hedwigia 39: 413. 1985 (1984)
Lophocolea bidentata
var. ß cuspidata Nees, Naturg. Eur.
Leberm. 2: 327. 1836; Lophocolea cuspidata (Nees) Limpricht; Chiloscyphus
cuspidatus (Nees) J. J. Engel
& R. M. Schuster Plants very loosely growing over
substratum and prostrate to ascending, very pale and hyaline green, 2.5--4 mm
wide when well developed, smaller phases 2.6--3 mm wide. Branching
frequent, the branches of Frullania-
and lateral intercalary types. Stems with cortical cells moderately differentiated, in 1(--2) layers of somewhat
smaller cells, the cortical and medullary cells thin-walled. Rhizoids
in tight fascicles from stem at bases of underleaves. Leaves
strongly succubous, the insertion distinctly recurved at ventral end, the
lines of insertion extending to stem midline dorsally; leaves alternate,
horizontal, widely spreading, broadly and rather evenly convex, a- to
sub-symmetrically ovate-trapezoidal to trapezoid, occasionally
subsymmetrically ovate, widest near base; apex 2-fid to 0.25--0.3; lobes
subcaudate, +/- parallel to diverging, the ventral lobe often larger,
tapering from a narrowly to broadly acute basal sector to a zone of up to 7
tiers of 2 laterally juxtaposed cells and, ultimately, terminating in a
piliform tip consisting of a uniseriate row of 5--10(--13) cells that are
variable but usually longer than wide, the lobe margins entire; dorsal margin
plane to deflexed, entire, often long decurrent; ventral margin entire or
sporadically with 1(--2) ciliiform projections. Cells
of leaf with very thin walls, with trigones minute to small and
concave-sided, the median cells nearly isodiametric, (27--)32--36(--40) /um
wide, 35--45 /um long; surface smooth.
Oil-bodies hyaline, 3--7
per cell, appearing coarsely papillose, broadly fusiform to ellipsoidal,
averaging 4 x 6 /um to 5 x 8--10 /um. Underleaves 1.6--2.7X stem width,
typically free and in close proximity to side nearest adjacent ventral base
of leaf, occasionally narrowly connate on 1 side, the insertion inverted
U-shaped, the underleaves plane to slightly
concave (ventral view), obtrapezoid to occasionally subrectangular, 2-fid
nearly to base (delimiting a lamina of a few cells high), the lobes
distinctly caudate, gradually tapering to a piliform tip normally consisting
of a uniseriate row of (4--)5--10, the lobe margins entire or frequently with
a few, often opposing ciliiform projections; lamina with a single process
variable in stature and position:
lobe-like to more often laciniiform to ciliiform, always smaller than
the main pair of lobes, the lamina margins sporadically armed with a few
accessory cilia. Specialized sexual reproduction lacking. Sexual
condition autoicous; plants nearly always fertile. Androecia
nearly always on lateral-intercalary branches, subfolious to subspicate;
bracts much smaller than leaves, the apex similar to leaves, except the lobes
acuminate to subcaudate; lobule margin involute, armed with an ill-defined,
low and rounded to tooth-like or distinct and ciliiform to laciniiform distal
projection, the margin otherwise with several cilia, the margin otherwise
entire and devoid of slime papilla or crenate-dentate; antheridia solitary,
the stalk uniseriate. Gynoecia terminal, mostly on main
shoot or lateral-intercalary branches; bracts much larger than leaves, 2-fid
to 0.35, the lobes narrowly attenuate, caudate distally, the margins strongly
reflexed, the dorsal margin entire or with a cilium, the ventral margin with
a few teeth or cilia; bracteole narrowly connate on both sides, 2-fid to
0.35--0.5. Perianth long and narrowly oblong, sharply trigonous and
prismatic throughout, the mouth 3-lobed, the lobes free for ca. 0.3--0.35 the
perianth length, each lobe conspicuously 2-fid, the segments acuminate to
(more often) subcaudate to caudate, entire or with 1--2 spinose teeth or
cilia, the lobes otherwise dentate-ciliate in basal half; keels sharp, with
wings common, often sparingly dentate-ciliate. Capsule
wide ovoid to ellipsoid, the walls 4-stratose. Spores
12.4--13.5 /um, with dense, low but sharply defined, close papillae and
short, simple or sometimes furcate, vermiculate markings. Elaters
tortuous, 7.2--8.6 /um wide. On wet
rocks, near running water, with other bryophytes also associated with damp or
inundated rock faces; also on thin soil and wood; low elevations; B.C.;
Alaska, Ark., Calif., Conn., Del?, Ga., Idaho, Ill., Ind., Md.?, Mass.,
Missouri, Mont., New Jersey, N.Y., N.C., Oreg., S.C., Tenn., Fa., Wash.,
W.Va.; Mexico; Cuba; South America (Venezuela) Europe; Asia; Africa
(including Reunion and Madagascar); Atlantic Islands; Pacific Islands
(including New Zealand); Australia. 4. Chiloscyphus apalachicolus
(R. M. Schuster)
J. J. Engel & R. M. Schuster, Nova Hedw. 39: 410. 1984 [1985] E Lophocolea apalachicola R. M. Schuster, Hepat. Anthoc.
N. Amer. 4: 195, pl. 521, figs. 1--12. 1980 Plants prostrate, light to whitish green, nearly pellucid, 2--2.8
mm wide. Branching irregular and sparingly, the branches of the Frullania- and ventral-intercalary
types. Stems with cortical cells not differentiated, the cortical and
medullary cells thin walled. Rhizoids from stem at base of
underleaves but becoming somewhat scattered.
Leaves strongly
succubous, the insertion nearly
longitudinal, somewhat oblique, not extending to stem midline dorsally and
delimiting a leaf-free strip 1--2 cells wide; leaves alternate, widely
spreading, strongly convex, often with an adaxial concavity at ventral base,
ovate; apex consistently 2-fid to 0.3--0.4, occasionally 3-fid; lobes usually
connivent, unequal, the ventral margin usually larger and little longer, long
acute, terminating in a uniseriate row of 3--5 +/- moniliform cells; dorsal
margin +/- decurved, a moderately well-defined cnemis often formed, the
margin sometimes with an accessory lobe, the ventral margin occasionally with
a lacinium; margins of lamina and lobes strongly and distinctly crenulate by
leptodermous, gibbous, protruding cells.
Cells of leaf leptodermous,
clearly with cells (as seen in profile) gibbously projecting on both
surfaces, the median cells 27--36(--40) /um wide, 35--45 /um long; surface
smooth. Oil-bodies unknown. Underleaves ca. 2X stem width,
distant, free from lateral leaves, the insertion crescent shaped, obovate to
obdeltoid, 2-lobed to 0.65--0.85; lobes lanceolate-acuminate; lamina margins
on each side with a ciliiform to laciniiform process. Asexual
Reproduction absent. Asexual Condition dioicous. Androecia
unknown. Gynoecia known only immature. Restricted
to humid and damp areas, over calcareous substrates, in partially evergreen
forests; Ala., Fla. Calcium-rich
environments; low elevations; Ala., Fla. Chiloscyphus apalachicolus is endemic to a small area in
western Florida into Alabama, found with Torreya
taxifolia and Taxus floridana,
and associated with Lejeuenea calcicola,
L. bermudiana, Dumortiera hirsuta,
Cololejeunea ornata, and Plagiochila
spp. This species is similar to Chiloscyphus latifolius and C. cuspidatus, but differs in having
the leaf cells gibbously projecting on both surfaces, lending the leaf and
underleaf margins distinctly crenulate.
The +/- moniliform uniseriate row of leaf lobe cells is also
distinctive. 5. Chiloscyphus appalachianus (R. M. Schuster) J. J. Engel
& R. M. Schuster, Nova Hedwigia 39: 410. 1984 [1985]. E Lophocolea appalachiana R. M. Schuster, Hep. Anthoc. N.
Amer. 4: 208, figs. 524, 525. 1980. Plants prostrate, in thin shiny patches, bright to deep green.
1.8--2.4(--2.8) mm wide. Branches strictly of
lateral-intercalary type. Stems with cortex in 1(--2) rows of
smaller, moderately thick-walled, in part pigmented cells; medullary cells
thin walled. Rhizoids sparsely developed, in tight fascicles from stem at
immediate base of underleaves. Leaves polymorphous, densely
chlorophyllose, strongly succubous, the insertion not recurved at ventral
end, the lines of insertion extending to stem midline dorsally; leaves
alternate, widely spreading, free dorsally, gently convex, +/- symmetrically
suborbicular to ovate to feebly oblate, rather evenly narrowed to the apex;
apex subtruncate, variable, undivided and entire (often) or 1-dentate to 2-dentate
to weakly 2-lobed; lobes (when distinct) merely acute or subacute, rarely
ending in a tooth with a uniseriate row of 2--3 cells; sinus crescentic;
dorsal and ventral margins broadly arched, entire. Cells
of leaf with walls thin, with trigones lacking, the median cells 25--34 /um
wide x 30--46 /um long; surface smooth.
Oil-bodies usually 2--4
(sometimes 6--16) per median cell, finely segmented and appearing papillose,
ovoid to ellipsoid. Underleaves ca 0.6--1X stem width,
distant, very narrowly connate on 1 or both sides, the insertion crescent- to
shallowly U-shaped, the underleaves 2-lobed to ca. 6 or more; lobes acute to
long acuminate; lamina margins entire, sometimes with a sharp lateral tooth
on 1 or both sides. Specialized sexual reproduction
absent. Sexual condition autoicous, often fertile. Androecia
on main shoot or elongated leafy branches, terminal but becoming
intercalary in position, subfoliose to slenderly spicate; bracts in 4--10
pairs, smaller than leaves, the apex evenly rounded, rarely truncate or
emarginate; antheridial stalk uniseriate.
Gynoecia on long leafy
shoots, or more often on abbreviated lateral branches with 1--4 gyres of
progressively larger leaves; bracts ovate to ovate-elliptical, the apex 2-dentate
or 2-dentulate, often subentire, the teeth small and often obscure, the
margins usually strongly reflexed, evenly arched, edentate; bracteole usually
considerably smaller than bracts, ovate, the apex usually obliquely 2-dentate
or 2-dentulate, but with 1 tooth often suppressed. Perianth
sharply trigonous, the mouth distinctly 3-lobed, each lobe shallowly 2-lobed,
the segments acute to subacute, the lobes with a few accessory teeth below
level of segments; keels strongly winged, the wings entire or coarsely
dentate to laciniate. Capsule not seen. Spores
15--19 /um, yellowish brown, delicately papillose. Elaters
2-spiral, yellowish brown, 8.5--10 /um wide. On
shaded, wet, noncalcaerous rocks; moderate to high elevations; N.C., Tenn. Chiloscyphus appalachianus is a polymorphic species most
similar to C. cuspidatus, but can
be distinguished by the deep green color, the smaller cell size, and the
subtruncate variable, undivided and entire or 1-dentate to 2-dentate to
weakly 2-lobed leaf (and male and female bract) apices, versus the pale green
color, larger cell size and sharply 2-fid leaves (and male and female bracts)
of C. cuspidatus. 6. Chiloscyphus profundus (Nees) J. J. Engel & R. M. Schust.,
Nova Hedw. 39: 421. 1984 [1985] Jungermannia profunda Nees, Naturg. Eur. Leberm. 2:
346. 1836; J. heterophylla Schrader; Lophocolea
heterophylla
(Schrader) Dumortier Plants exceedingly polymorphic, creeping, pale to yellow green,
well developed shoots 1.2--1.6(--2) /um wide, but often smaller, 1--3 cm
long. Branching common, the branches of the lateral- and
ventral-intercalary types. Stems poorly undifferentiated, the
cortical cells in a single layer of slightly thick-walled cells; medullary
cells thin walled. Rhizoids frequent, from stem at bases
of underleaves. Leaves
strongly succubous, the insertion
nearly horizontal, not extending to stem midline dorsally and delimiting a
leaf-free strip 1 cell wide; leaves strongly alternate, widely spreading,
weakly convex, oblong to oblong-quadrate, the dorsal and ventral margins
entire, often nearly straight, nearly parallel, the leaves strongly
polymorphous, the lower leaves 2-lobed on mature shoots, the lobes obtuse or
acute, terminated in a single cell, the lamina margins not decurrent, the
upper leaves larger, the apex rounded-truncate, retuse to broadly and
shallowly emarginate, the lobes low and rounded when present; sinus obtuse to
rectangulate; leaves almost parallel-sided and often slightly narrowed near base,
the dorsal margin short-decurrent; leaves undivided in subsp. cladogyna. Cells
of leaf with walls thin, with trigones minute to occasionally slightly
bulging (in sun forms), the median cells (20)23--30 /um wide, 28--35 /um
long; surface smooth. Oil-bodies 4--8(--10) per cell,
obviously segmented, ovoid to ovoid-fusiform, rarely spherical, 2--4 x 5--8 /um
to 4 x 5.5--12 /um, or 4.5 x 9.5 /um. Underleaves ca. as wide as stem, free
from lateral leaves, the insertion crescent- to shallowly U-shaped, subrectangular
to obdeltoid, 2-lobed to ca. 0.7--0.9; lobes lanceolate-acuminate, often with
a sharp lateral tooth on 1 or both sides toward base; lamina margins often
with a dentiform to laciniiform process on 1 or both sides. Specialized
asexual reproduction rare, by 1--2 celled gemmae or linear multicellular
bodies. Sexual condition paroicous, rarely heteroecious. Androecial
bracts in 2--4(--5) pairs, erect, slightly smaller to slightly larger
than vegetative leaves, saccate at base, the dorsal margin with an incurved
tooth; antheridial stalk uniseriate. Gynoecia terminal on leafy branches,
the branches often relatively abbreviated; bracts oblong to oblong-quadrate,
the apex retuse or 2(rarely 3--4)-lobed; bracteole 2-fid, the lobes
lanceolate, acuminate, the lamina margins with a few teeth. Perianth
trigonous to the middle or below, terete near base, the mouth shallowly 3-lobed,
the lobes coarsely and irregularly dentate or lobulate; keels usually lacking
wings. Capsule elliptic, the wall of 4--5 layers. Spores
8--11(--13) /um, yellowish brown, nearly smooth. Elaters
equal in diameter to spores. On
organic substrates, moist decaying wood, occasionally moist mineral soil; low
to high elevations; B.C., Ont., Que.; Ark., Calif., Ill., Iowa, Kans., Ky., La.,
Maine, Md., Mich., Miss., Mo., Mont., N.H., N.C., N.Dak., Ohio, Okla., Oreg.,
Pa., S.Dak., Tenn., Va., Wash., Wis.; Eurasia; n Africa. 1. Leaves, at least on weak shoots, and in the
basal 0.5 of mature, fertile shoots sharply 2-dentate; gemmae normally
absent; gynoecia on elongate, leafy shoots, with ♀ bracts large (as long or
longer than leaves)
............................ 6a. Chiloscyphus
profundus subsp. profundus 1.
Leaves, even on small, weak shoots, and usually throughout mature, fertile shoots,
rounded-oblong, entire, with apex truncate; gemmae usually freely developed;
gynoecia, at least in part, on abbreviated, leafless lateral shoots, with ♀
bracts smaller than leaves
.............................. 6b. Chiloscyphus
profundus subsp. cladogyna 6a. Chiloscyphus profundus (Nees) J. J. Engel & R. M. Schuster
subsp. profundus Leaves, at least on weak shoots, and in
the basal 0.5 of mature, fertile
shoots sharply 2-dentate; gemmae
normally absent; gynoecia on
elongate, leafy shoots, with ♀ bracts large (as long or longer than
leaves). 6b. Chiloscyphus profundus (Nees) J. J. Engel & R. M. Schuster
subsp. cladogyna R.M.
Schuster E Chiloscyphus profundus subsp. cladogyna R.M. Schust., Hep. Anthoc. No. Amer. 4: 223. 1980. Leaves, even on small, weak shoots, and
usually throughout mature, fertile
shoots rounded-oblong, entire, with apex truncate; gemmae usually freely developed; gynoecia, at least in part, on abbreviated, leafless lateral
shoots, with ♀
bracts smaller than leaves. Evergreen
or semi-evergreen coastal swamps, often on tree bases, also coastal sand
dunes on bases of Quercus virginiana;
low elevations; Fla., Miss., N.C., S.C.
7. Chiloscyphus minor (Nees) J. J. Engel & R. M. Schuster,
Nova Hedw. 39: 419. 1985 (1984)
Lophocolea minor
Nees, Naturg. Eur. Leberm. 2: 330.
1836 Plants rather delicate, with a pungent odor, prostrate, yellow
green, quite small, usually only 0.5--1(--1.2) mm wide, 12--18(--15) mm
long. Branching frequent,
the branches of the Frullania- and
lateral-intercalary types. Stems undifferentiated, the cells
leptodermous. Rhizoids rather frequent, from stem at bases of
underleaves. Leaves strongly succubous, the
insertion strongly oblique, not extending to stem midline dorsally and
delimiting a leaf-free strip 1--3 cells wide; leaves strongly alternate,
widely spreading, slightly to moderately convex, oblong-ovate to subquadrate;
apex 2-lobed to 0.25--0.35, the lobes bluntly acute to obtuse, but with
gemmae formation often the entire distal sector of the leaf erose and
the lobes becoming obsolete, the gemmiparous leaves becoming abnormal and
erose-margined although not reduced in size; sinus usually rectanulate
to obtuse or lunate, rarely acute; dorsal and ventral margins often
nearly straight and nearly parallel, the dorsal margin long decurrent. Cells
of leaf thin walled, with trigones minute or lacking, the median cells
20--25(--30) /um; surface smooth. Oil-bodies 3--10 per cell, segmented,
subspheric to ovoid, 3--4 to 4--5 x 6--7 /um. Underleaves
about as wide as stem to slightly wider, distant, free from lateral leaves,
the insertion crescent- to shallowly U-shaped, the underleaves subrectangular
to obdeltoid, 2-lobed to 55--0.75; lobes lanceolate to linear-lanceolate;
lamina margins often with a sharp tooth on 1 or both sides. Specialized
asexual reproduction by gemmae, the gemmae universally developed on
leaves, usually abundantly produced, also from underleaf lobes, female bracts
and perianth mouth; gemmae yellow green, initially spherical and 1-celled,
but eventually usually forming irregular 2-many celled masses or short
filaments. Sexual condition dioicous or paricous. Androecia
when paricous with male bracts usually with vestigial or aborted antheridia;
when dioicous the male bracts supposedly saccate at base of dorsal
margins. Gynoecia on long leafy shoots; bracts usually narrowly oblong to
cuneate-oblong, the lobes blunt to rounded, sometimes erose with gemmae
formation; bracteole at least 0.5 the bract length, 2--3-lobed. Perianth
narrowly elongate, sharply triquetrous above, the mouth 3-lobed, the lobes
coarsely toothed, occasionally gemmiparous; keels lacking wings or at most an
isolated low wing present. Capsule ellipsoidal. Spores
ca. 10 /um, nearly smooth. Elaters 8--10 /um wide, 2-spiral. Basic
substrates, dry loamy or sandy calcareous banks, also decaying logs; low to
high elevations; Alta., B.C., Man., N.B., Nfld. (Nfld., Labr.), N.W.T., N.S.,
Ont., Que., Yukon; Alaska, Colo., Conn., Idaho, Ill., Ind., Iowa, Kans.,
Maine, Mass., Mich., Minn., N.J., N.Mex., N.Y., Penn., Utah, Vt., Va., W.Va.,
Wis.; Eurasia. Chiloscyphus minor is found in a wide diversity of
sites, but mostly restricted to basic regions. 8.
Chiloscyphus muricatus
(Lehmann) J. J. Engel & R. M. Schust. Nova Hedw. 39: 419. 1984 [1985] Jungermannia muricata
Lehmann, Linnaea 4: 363. 1829. Lophocolea
muricata (Lehmann) Nees Plants delicate, wooly in appearance, soft, loosely prostrate to closely
adnate, pale green to yellow green to pale whitish green. small, to 1 mm wide. Branching common, the branches of the
Frullania- and ventral-intercalary
types. Stems with branching common, the branches of Frullania- and ventral-intercalary types; stems smooth, the
cortex only slightly differentiated by smaller cells on ventral side, both
cortical and medullary cells with moderately thick walls. Rhizoids
from stem in an ill-defined zone at base of underleaves.
Leaves succubous to subransverse, with insertion faintly recurved
at ventral end, the lines of insertion extending to stem midline dorsally;
leaves with dorsal surface densely hispid except near ventral base, armed on
dorsal surface with erect, 1--3 celled echinae which are juxtaposed 1 each
over the lumen of most cells, the ventral surface with ventral half armed
with 2--4-celled scattered, erect echinae; leaves varying from clearly
alternate to subopposite, subhorizontal, broadly convex, the apices broadly
deflexed and oriented at ca. 90 º with remainder of leaf, the leaves
subsymmetrically broad ovate; apex 2-fid to 0.2, the lobes acute, the margins
with opposing 1--2 celled sharp teeth; lamina margins spinose-dentate to the
base. Cells of leaves with walls moderately thickened, with trigones
small to medium and then the sides straight, median leaf cells without spines
14--18 /um wide, 17--19 /um long, the spine-bearing cells about the same
size. Oil-bodies large for cell size, pellucid to greyish, 2--3 per
cell, papillose, spherical and (2--)3--4 /um to ovoid or ellipsoid and
(2.5--)3--3.5 x 4--5.5 /um. Underleaves narrowly connate on 1
side, occasionally free, 2-lobed to 0.75--0.8 or deeper, distant, cuneate to
long rectangular, 1.4--1.6X stem width; lobes long-linear to subulate, the
margins with cilia that are often branched, the surface often with scattered
ciliiform processes. Specialized asexual reproduction lacking.
Sexual condition autoicous. Androecia
terminal or becoming intercalary in position on leading shoots or
branches, short spicate; bracts much smaller than leaves, the surface and
margins with armature similar to leaves; antheridia solitary, the stalk
uniseriate. Gynoecia terminal on main shoots and ventral-intercalary
branches; bracts much larger than leaves, with surface armature similar to
leaves, the apex short 2-fid, the lamina margins spinose-dentate to
dentate-ciliate; bracteole much smaller than bracts, connate on one
side. Perianth long exserted, cylindrical or dorsiventrally compressed,
narrowly oblong to clavate, the surface densely hispid; perianth not to
slightly expanding toward the wide, distinctly 3-lobed mouth; lobes shallowly
2-fid, the segments medium acute to subacuminate, with margins spinose
dentate as in leaves, the lobes otherwise lobulate and spinose dentate. Seta
with 10 rows of outer cells surrounding an inner core of 9 scattered, much
smaller cells. Capsule wide ellipsoid, the wall 2-stratose (and then 17--18 /um
thick), or 3-stratose (and then 20--22 /um); outer layer of cells with
nodular to feebly spine-like thickenings, the thickenings mostly on
longitudinal walls and only sporadically on transverse walls; innermost layer
of cells with semiannular bands common.
Spores 10.1--12.4 /um, pale
brown, with faint, dense, low, close, papillose and sporadic
short-vermiculate markings. Elaters rather tortuous, 7.2--9.6 /um
wide. On rocks, bases of trees and
decaying logs, among other liverworts and mosses; low to high elevations; Ga., N.C., S.C.; Mexico; West Indies;
Central America (Honduras); South America; Asia (Indonesia); Africa; Australia;
Pacific Islands (New Zealand, Papua New Guinea, Fiji). 9. Chiloscyphus polyanthos (Linneaus) Corda in Opiz,
Beiräge zur Naturg. 12: 651. 1829 Jungermmania polyanthos Linneaus, Spec. Pl. 1: 1131.
1753; Chiloscyphus rivularis
(Schrader) Hazsliln; Jungermannia
pallescens var. rivularis Schrader;
Marsupella polyanthos (Linneaus)
Dumortier Plants loosely creeping to weakly ascending, pure to dull green
to yellow-green. usually 2.4--3.5(--4.2) mm wide x 2--5(--6) cm long. Branching
frequent, the branches of the Frullania-
and lateral-intercalary types. Stems with cortical cells in 1--2
rows of smaller cells. Rhizoids frequent, from stem at bases
of underleaves. Leaves strongly succubous, the insertion nearly longitudinal, not
extending to stem midline dorsally and delimiting a leaf-free strip 1--2
cells wide; leaves strongly alternate, widely spreading, slightly to
moderately convex, rounded quadrate to very short-rectangulate; apex
undivided, occasionally retuse, occasionally emarginate (especially on lower
leaves), rounded or truncate; dorsal and ventral margins subequally
arched, or the ventral a little more strongly so, entire, the dorsal margin
distinctly decurrent; Cells of
leaf leptodermous, with trigones absent, the cells relatively small, the median
cells usually 21--28 /um wide x 28--35(--40) /um long, rarely to 30--35 x
35--40 /um. Oil-bodies usually 1--2, occasionally 3--4 per cell, distinctly
but finely segmented, fusiform to oval, from 4--5 x 6.512 /um up to 7 x 9--12
/um, isolated oil-bodies larger. Underleaves as wide as stem or
narrower, free from leaves, the insertion crescent-shaped, the underleaves
elliptic to subrectangular, 2-lobed to 0.25--0.5; lobes narrowly acuminate to
narrowly attenuate, at times with a tooth toward the base, lamina margins
with a tooth on one or both sides. Asexual reproduction unknown. Sexual condition autoicous. Androecia
on leading shoots; bracts in several pairs, leaflike, with a small inflexed
lobule at dorsal base; antheridia 1 per bract, the stalk uniseriate. Gynoecia
on abbreviated lateral-intercalary branches; bracts smaller than vegetative
leaves, 2-lobed, often unequally so; bracteole similar to bracts. Perianth
cylindrical below, inflated-trigonous above, the mouth wide, 3-lobed, the
lobes entire or with a few small teeth.
Calyptra long emergent from
beyond perianth. Capsule wall 4-stratose. Spores 12--18 /um, yellowish brown,
finely granular. Elaters 2-spiral, 7--8.5 /um On humus
and soil over rock at the edges of streams and ponds, often in shaded or
semishaded sites; low to high elevations;
Holarctic in North America, Eurasia. 1.
Plants pure to dull green to yellow green; leaves contiguous to weakly
imbricate; median leaf cells usually 21--28 /um wide
................................. 9a. Chiloscyphus
polyanthos var. polyanthos 1.
Plants deep green to nearly blackish green; leaves remote to contiguous;
median leaf cells smaller, 19--24 u wide ..................................
9b. Chiloscyphus polyanthus var. rivularis 9a. Chiloscyphus polyanthos (L.) Corda var. polyanthos Plants
pure to dull green to yellow green; leaves contiguous to weakly imbricate;
median leaf cells usually 21--28 /um wide. On humus
and soil over rock at the edges of streams and ponds, often in shaded or
semishaded sites, lacking from strongly calcareous sites; low to high elevations;
Miquelon I.; Alta., B.C., Nfld. &
Labr. (Nfdl.), Man., N.S., Ont., Que.; Alaska, Calif., Colo., Conn., Idaho, Ill.,
Ind., Iowa, Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nev., N.H., N.Mex.,
N.Y., N.C., Ohio, Oreg., Pa., S.C., Tenn., Tex., Utah, Vt., Va., Wash., Wis.,
Wyo.; Mexico; Eurasia. 9b. Chiloscyphus polyanthos (L.) Corda var. rivularis (Schrad.) Lindenb. &
Arnell, Jungermannia pallescens var. rivularis Schrad., Syst. Samml. Crypt. Gew. 2: 7. 1797.
Chiloscyphus polyanthos var. rivularis (Schrad.) Lindenb. & Arnell, Kongl. Svenska
Vetensk.-Akad. Handl. (n. ser.) 23 (5): 24.
1889. Plants
deep green to nearly blackish green; leaves remote to contiguous; median leaf
cells smaller, 19--24 u wide. Found
submerged in creeks, although becoming exposed with drying of the niche; low
to high elevations; Alta., N.B., N.S.,
Ont.; Ark., Calif., Colo., Del., Maine, Mass., Mich., Mo., Nev., N.Y., Oreg., Pa., Wash., Wyo.; Eurasia. Chiloscyphus polyanthos var. rivularis is likely to occur throughout the range of the species;
It is considered to be the aquatic form of the species that is present in
standing, peaty waters such as ditches and ponds. 10. Chiloscyphus pallescens
(Ehrhart ex
Hoffmann) Dumortier, Syll. Jungerm. Eur., 67. 1831 Jungermannia pallescens Ehrhart ex Hoffmann, Deutschl.
Fl. 2: 87. 1795 (1796); Chiloscyphus
polyanthos var. pallescens
(Ehrhart) C. Hartman; Jungermannia polyanthos var. pallescens (Ehrhart) Lindenberg Plants prostrate to decumbent, pale whitish or yellowish green,
rather translucent, the shoots usually 1.8--3.2(--3.5) mm wide. Branching
sparse and irregular, the branches of the Frullania
type. Stems with cortical cells in 1--2 rows of smaller cells. Rhizoids
few, from stem at bases of underleaves.
Leaves with insertion
almost longitudinal, the lines of insertion not extending to stem midline
dorsally and delimiting a leaf-free strip several cells wide; leaves
alternate, widely spreading, somewhat convex, oblong-quadrate, rarely
rounded-quadrate to +/- lingulate; apices undivided, rounded-truncate to
truncate, occasionally shallowly retuse, the leaves at base of stem typically
2-lobed by acute to obtuse lobes; margins entire. Cells
of leaf walls thin and delicate, with trigones lacking, the cells large, the
median cells 33—40(--42) /um wide x 40--50 /um long; surface smooth. Oil-bodies
relatively small, (3--)4--6(10--12) to (6--)8--12(--15) per cell, finely but
distinctly segmented, ellipsoidal to fusiform or ovate, from 4--5.5 x 8--9 /um
up to 5 x 10--11 /um. Underleaves as wide as stem or
somewhat narrower, distant, free from leaves, the insertion weakly crescent
shaped, the underleaves subrectangular, 2-fid to 0.5--0.8; lobes subulate to filiform;
lamina margins on each side with 1(--2) short to filiform teeth. Specialized
asexual reproduction absent. Sexual condition autoicous. Androecia
intercalary in position on main shoots; bracts in 4--6 pairs, not
differentiated from leaves, aside from a development of a small 1--2-dentate
lobule; antheridial stalk uniseriate.
Gynoecia on abbreviated
lateral-intercalary branches; bracts much smaller than vegetative leaves,
shallowly 2-lobed; bracteole ovate-lanceolate. Perianth
stoutly cylindrical-subclavate, the mouth rather deeply 3-lobed, the lobes
sharply dentate or ciliate, occasionally spinose-dentate or subentire. Calyptra
slightly to distinctly exserted beyond perianth at maturity. Capsule
wall 5(--6)-stratose. Spores yellowish brown, 13--15(--19)
µum in diameter, finely granulose. Elaters 8--9 /um wide. Often
near running or standing water, moist soil or rock or on organic substrates;
low to high elevations; North America, Eurasia. Chiloscyphus pallescens is a Holarctic species, common in
calcareous sites, transcontinental in North America and widespread in Europe. 1.
Plants translucent, the shoots smaller, 1.8--3.2 mm wide; leaves usually
oblong-quadrate to +/- lingulate, 1--1.4X longer than wide; apex rounded-truncate
to truncate, occasionally shallowly retuse .......................... 10a. Chiloscyphus pallescens var. pallescens 1. Plants relatively opaque, the shoots
larger, 3.5--5 mm wide; leaves rounded quadrate, usually as broad as long, at
times wider than long; apex usually rounded, exceptionally truncate-retuse
..................................... 10b. Chiloscyphus pallescens var. fragilis 10a. Chiloscyphus pallescens
(Ehrhart)
Dumort. var. pallescens Plants
translucent, the shoots smaller, 1.8--3.2 mm wide; leaves usually
oblong-quadrate to +/- lingulate, 1--1.4X longer than wide; apex
rounded-truncate to truncate, occasionally shallowly retuse. Often
near running water nearly always above flood level or near standing water,
usually partly or entirely in shade, moist soil or rock or on organic
substrates such as decaying logs and humus; low to high elevations; B.C., Nf.d.
& Labr. (Nfld., Labr.), N.W.T., N.S., Ont., Que., Yukon; Ala., Alaska, Ark.,
Calif., Colo., Conn., Fld., Idaho, Ill., Ind., Iowa, Kans., Ky., Maine, Mass.,
Mich., Minn., Mo., Neb., N.H., Nev., N. Mex., N.Y., N.C., Ohio, Oreg., Pa., R.I.,
S.Dak.,Tenn., Utah, Vt., Va., W.Va., Wyo.; Eurasia. 10b. Chiloscyphus pallescens (Ehrhart) Dumort. var. fragilis (Roth) Müll. Frib. ex R. M.
Schusteer, Amer. Midl. Naturaalist 42: 587. 1949 Jungermannia fragilis Roth, Tent. Fl. Germ. 3:
370. 1800; Chiloscyphus fragilis (Roth) Schiffner; C. pallescens var. fragilis
(Roth) Müll. Frib.; C. polyanthos var.
fragilis (Roth) Müll. Frib. Plants
relatively opaque, the shoots rather large, 3.5--5 mm wide; leaves rounded
quadrate, usually as broad as long, at times wider than long; apex usually
rounded, exceptionally truncate-retuse.
Slow-moving
streams, shallow pools and meadows subject to inundation, at times on
dripping rocks or on soil over rocks at the margins of brooks or in stream
beds; low to high elevations; B.C., N.B., Nfld. & Labr. (Nfld., Labr.),
N.S., Ont., P.E.I., Que.; Alaska, Calif., Colo., Conn., Idaho, Ind., Iowa,
Kans., Maine, Mass., Mich., Minn., Mo., Mont., N.H., N.J., N.Y., N.Mex., N.C.,
Oreg., Pa., Tenn., Vt., Va. Utah, Wash., W.Va., Wis., Wyo.; Mexico
(Veracruz); Eurasia. Chiloscyphus pallescens var. fragilis occurs sporadically throughout the range of the species,
but drops out southward. 3. CRYPTOLOPHOCOLEA L. Söderström, Crandall-Stotler, Stotler & Váňa, Phytotaxa
97(2): 39. 2013 * {Greek kryptos, hidden, lophos, crest, and koleos,
sheath, alluding to similarity with Lophocolea] Laura Briscoe John J. Engel Chiloscyphus subg. Connati
(Lindenberg) J. J. Engel Plants
loosely prostrate to ascending to erect, light green to yellow green to
whitish green, at times becoming pale brown or yellow-brown in older sectors,
to 5 mm wide. Stem with cortex
poorly to at most moderately differentiated, in 1--2(--3) layers of rather
thin- or thick-walled cells typically slightly to distinctly smaller than
those of the medulla; branching common, lateral-intercalary [terminal]. Rhizoids colorless, from stem at base of underleaf
[rarely becoming scattered]. Leaves
with insertion strongly succubous, non-recurved at ventral end, the lines of
insertion [extending to stem midline dorsally or] not extending to the stem
midline; leaves subopposite to opposite, free or connate dorsally, plane or
convex, the apex short-2-fid [or undivided], the ventral lobe larger; lobe
margins entire; lamina margins entire or with several dentiform to
laciniiform teeth. Median cells
thin-walled, trigones [medium to knot-like to nodulose or] absent to minute;
surface smooth or finely papillose [tuberculate]. Underleaves
connate on both sides. Sexual
condition dioicous or autoicous. Androecia
on main shoots or leading lateral-intercalary or Frullania type
branches; bracts considerably smaller than leaves, the
dorsal base dilated and forming an infolded flap or lobule; lobule margin
crenate or dentate, at times with a few ciliiform projections, at times with
a several-celled tooth in distal sector; antheridia 1 per bract, the stalk
biseriate. Androecia on main shoots or leading lateral-intercalary or Frullania
type branches; bracts considerably smaller than leaves, the dorsal base dilated and forming an infolded
flap or lobule; lobule margin crenate or dentate, at times with a few
ciliiform projections, at times with a several-celled tooth in distal sector;
antheridia 1 per bract, the stalk biseriate. Gynoecia on main shoot or lateral-intercalary branches. Perianth
sharply trigonous and at times prismatic throughout, the mouth shallowly to
more often deeply 3-lobed, the lobes subequal, typically distinctly 2-fid [in
a few species undivided merely dentate-laciniate], the segments narrowly
acute to acuminate to caudate, entire margined, the lobes otherwise coarsely
dentate to laciniate below level of segments; keels with wings common, at
times conspicuously high, at times low and incomplete; perigynium absent. Capsule elliptic to long
ellipsoid-cylindrical, walls 4--6(--7) stratose. Elaters weakly to distinctly tortuous, 2-spiral. Spores with a low, close network of
anastomosing vermiform ridges delimiting irregular areolae [variously finely
papillose]. Species ca. 29 (1 in the flora): North America, West Indies, Central
America, South America, Asia, Africa, Pacific Islands (including New
Zealand), Australia. The genus Cryptolophocolea
was recently described to accommodate species formerly recognized in Chiloscyphus subgen. Connati. As a group, species in this genus have
several characters that set them apart from Chiloscyphus s. str., including perianths elongate, trigonous,
with winged keels. They also have distinct, large underleaves that are
connate with lateral leaves on both sides. The delineation of this genus is
supported by molecular evidence. The genus is pantropical and temperate
Australasian in range. SELECTED REFERENCES:
Söderström, L., B. Crandall-Stotler, R. E. Stotler, J. Váňa, A.
Hagborg, & M. von Konrat. 2013. Notes on early land plants today. 36.
Generic treatment of Lophocoleaceae (Marchantiophyta). Phytotaxa 97(2):
36--43. Söderström, L., B. Crandall-Stotler, R. E. Stotler, J. Váňa, A.
Hagborg, & M. von Konrat. 2013. Notes on early land plants today. 43. New
combinations in Lophocoleaceae (Marchantiophyta). Phytotaxa 112(1): 18--32. Engel, J. J.
2014. Studies on Lophocoleaceae XXIII. Novelties in Heteroscyphus Schiff. together with refinements in Cryptolophocolea L. Söderstr.,
Crand.-Stotl., Stotler & Váňa and Leptoscyphus
Mitt., Nova Hedw. 99(1--2): 157--170. 1. Cryptolophocolea martiana
(Nees) L.Söderstr., Crand.-Stotl. & Stotler, Phytotaxa 112: 20. 2013 Lophocolea
martiana Nees Syn. Hepat. 2:
152. 1845; Chiloscyphus jugalifolius
Taylor; C. martianus (Nees) J. J.
Engel & R. M. Schuster Plants to 2.5 mm
wide. Stem cortical cells
moderately differentiated in 1--2(--3) layers of somewhat smaller, weakly
thick-walled cells, the medullary cells thin-walled. Leaves with the lines
of insertion not extending to stem midline dorsally, with a leaf-free strip
of 1--3 stem cells wide; leaves subopposite,
nearly horizontal, widely spreading, free dorsally, weakly convex to nearly
flat, nearly symmetrical, narrowly ovate-triangular, the apex narrowed, 2-dentate
to shallowly 2-lobed to 0.15; lobes dentiform, the ventral sometimes larger,
acute to apiculate to blunt; sinus subtruncate to lunate; dorsal and ventral
margins nearly straight to slightly arched.
Median cells
(24--)35--43(47--50) x (20--)24--30(--33) /um, surface faintly papillose. Oil-bodies unknown. Underleaves ca. 1.2--2 x stem width,
the insertion inverted U-shaped, the underleaves distant to approximate,
concave (ventral view), stoutly subrectangular to obtrapezoid, 4(--5)-lobed
to 0.6-0.7 the length at median sinus, the lobes slender, narrowly
lanceolate, sporadically with an accessory tooth, the underleaf margins
otherwise entire. Specialized asexual
reproduction lacking. Sexual
condition autoicous. Androecia terminal
but becoming intercalary, subfolious to weakly spicate, with 6--12 pairs of
bracts; bracts smaller than leaves, the dorsal-basal portion ventricose, the
apices emarginated to 2-lobed.
Gynoecia with bracts larger than leaves, erect and ensheathing perianth,
2-fid to 0.3 the length, margins with 1--2(--3) teeth; perianth narrowly
oblong, sharply trigonous, sides sometimes concave, the mouth deeply 3-lobed, each lobe
distinctly 2-fid, segment acuminate, lobe margins dentate to laciniate. Capsule elliptic, the walls 4--5
stratose. Elaters somewhat
tortuous, 2-spiral, 7.2-8.2 /um wide, pale reddish brown. Spores 10.6--12 /um, the wall
thin. On moist humus at stream edges and at bases of trees
and shrubs; wet and swampy regions of se Coastal Plain; low elevations; Fla.,
La., Miss., Mexico, West Indies; Central America; South America (tropical). Cryptolophocolea martiana
is a tropical to subtropical species that is widespread in the New World and
can be found in North America only in mesic, frost-free zones of the
southeast. It can be distinguished from species of Chiloscyphus by the subopposite leaves, which are connate with
adjoining underleaves. [[For
illustrations see individual genera on BFNA]] |