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BFNA Title: Lembophyllaceae |
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78. LEMBOPHYLLACEAE Brotherus William
R. Buck Plants
medium-sized to large, pale to dark green or golden, terrestrial, epipetric
or epiphytic, often somewhat stipitate. Stems
irregularly pinnate to regularly 2- or 3-pinnate, terete-foliate; cortex of
small and incrassate cells; hyalodermis absent; pseudoparaphyllia foliose;
paraphyllia absent; axillary hairs with 1--2 short, brown basal cells and
1--4 elongate, hyaline apical cells. Stem
and branch leaves often differentiated, loosely appressed, mostly ovate
and concave; costa single or double and short, sometimes stipe and stem
leaves with a weak costa and branch leaves with a strong costa; median
laminal cells short to linear, smooth or prorulose; alar cells mostly well
differentiated, often excavate, mostly quadrate. Sexual condition dioicous. Seta
mostly long, sometimes short, smooth or roughened throughout or just
distally. Capsule cylindric,
mostly erect and symmetric, but sometimes curved and horizontal; annulus
typically differentiated; operculum conic to short-rostrate, straight;
peristome typically perfect but sometimes reduced, exostome when not reduced
cross-striolate on outer surface proximally, papillose distally, when reduced
variously striate, papillose or smooth, endostome with high or low basal
membrane, segments long and narrowly perforate, cilia nodose or
appendiculate. Calyptra cucullate, smooth or slightly
roughened, naked or hairy. Genera 14, species ca. 50 (3 genera, 7 species in the
flora): mostly south-temperate, but extending into the tropics and The bulk of genera and species of the Lembophyllaceae
occur in the Southern Hemisphere, with only a few genera in the flora region.
The plants tend to have terete-foliate stems with ovate, concave leaves. The
costa is variable, sometimes on different leaves of the same plant. Many of
the genera have perfect hypnoid peristomes, but there are genera (e.g., Bestia) with reduced peristomes. The calyptrae being frequently hairy is an unusual character
in the flora region. Selected References
Crum, H. A. 1987. Bestia, Tripterocladium, and Isothecium: an explication of
relationships. Bryologist 90: 40--42. Quandt, D., S. Huttunen, R. Tangney and
M. Stech. 2009. Back to the future? Molecules take us back to the 1925
classification of the Lembophyllaceae (Bryopsida). Systematic Botany 34:
443--454. 1. Costa mostly short and double, rarely single (isolated
leaves); branch and stem leaves well differentiated ……………………………………………………………….
3. Tripterocladium, p. XX 1. Costa consistently single; branch and stem leaves
similar. 2. Median laminal cells less than
4:1; capsules erect with reduced peristomes ……. 1. Bestia, p. XX 2. Median laminal cells more than
5:1; capsules often inclined with perfect peristomes … 2. Isothecium, p. XX 1. BESTIA Brotherus in A. Engler & K. A. E. Prantl,
Die natürlichen Pflanzenfamilien 1(3): 859. 1906 * [For George N. Best,
1846--1926, American bryologist] James
R. Shevock Plants
robust, forming large descending mats. Stems pinnate or irregularly 2-pinnate;
branches generally curved to one side when dry. Leaves of stems and branches similar, ovate, broadly acute,
somewhat concave at base, imbricate when dry but erect-spreading when moist;
margins serrulate in distal 1/5; costa single, stout, distally toothed
abaxially; laminal cells short, thick-walled, those in the distal portion of
the leaf projecting at their distal ends as prorulae. Specialized asexual reproduction lacking. Capsules cylindrical, erect
and symmetric; peristome double, well formed, somewhat reduced. Species 1: Western North America, endemic to coastal The genus Bestia
has had a rather confusing taxonomic history causing difficulty in
determining its family placement depending on which species
of Bestia was critically examined
and what combination of morphological characters were considered most
important for inferring relationships (J. R. Shevock et al. 2008). H. A. Crum
(1987) placed what was to become this unispecific genus in close relationship
to Isothecium, in the Brachytheciaceae. The other species, Bestia vancouveriensis
was transferred to the Thamnobryaceae as Porotrichum vancouveriense (Kindberg ex Macoun) H.A. Crum,
and subsequently elevated to generic rank as Bryolawtonia vancouveriensis.
Bestia is a robust, distinctive
species endemic to coastal Although Bestia
has been attributed to several families over the years, recent DNA studies
suggest that both Bestia and Isothecium are closely related and
should be transferred to the Lembophyllaceae. Bestia is morphologically similar in appearance to some species
of Isothecium (especially I.
myosuroides), but it has several gametophytic features that readily
separate it, primarily median cells uniformly short (less than 4:1) whereas the juxtacostal cells in Isothecium are elongate to 8:1. The
long, straight, cylindric capsules with smaller peristome teeth differ
markedly from the rather hypnaceous curved capsules in Isothecium. SELECTED REFERENCES Crum, H. A. 1987. Bestia, Tripterocladium,
and Isothecium: an explication of relationships. Bryologist 90:
40--42. Shevock, J. R, D. H. Norris and A. J. Shaw. 2008. Identification,
distribution and family placement of the pleurocarpous moss Bestia longipes (Sull.
& Lesq.) Broth. Madroño 55: 291--296. 1.
Bestia longipes
(Sullivant & Lesquereux) Brotherus in A. Engler & K. A. E. Prantl,
Nat. Pflanzenfam 1(3): 859. 1906 E F Alsia longipes Sullivant &
Lesqeureux, Musci Bot.-Amer. (ed. 2), 66. 1865; A.
sullivantii Lesquereux ex
Fleischer Plants green
to deep green, growing in rather pendulous wefts. Stems to 12 cm, pinnately branched with secondary branches to 1
cm; rhizoids rarely present, pale
brown, smooth and essentially unbranched, inserted on stem base; axillary
hairs 3-celled with 2 basal brown
cells, only 4--5 \um wide at insertion but 10 \um wide distally, with the
terminal cell hyaline and about 40 \um in length; pseudoparaphyllia about as
broad as long, serrate at base, short cells throughout. Leaves to 1.6 mm, 3--5:1, symmetric and straight; margins narrowly
recurved at least near mid leaf, decurrent at base; median cells smooth, 5--6
\um wide, 1.2--1.8:1, thick‑walled (lumen wall ratio 2--4: 1), not or
inconspicuously porose; basal, juxtacostal cells somewhat more elongate,
3--4: 1; alar cells quadrate, thick-walled, with rounded lumina, extending
more than 20 cells up the margins and extending inward to the basal plica,
auriculate at the small decurrency; cells of apical region similar to median
laminal cells, often prorate-spinose in a small patch near the costal apex;
costa extending to within 10 cells of the apex, distal portion with many
abaxial spines. Perigonia nearly spherical on a short stalk, bud‑like with bracts
broadly ovate, 1.2--1.4:1, mostly rather sparsely cloaking and inserted on
the ventral side of the main axes; perichaetia on ventral side of main erect
axes, the bracts 2--3 mm, ecostate or with a short costa, strongly convolute
around the seta; inner perichaetial bracts with post-fertilization
development abruptly acuminate from an elliptic base. Seta
straw-colored, rarely reddish tinged, to 1.5 cm, erect, sporophyte often
shorter than adjacent lateral branches.
Capsule pale brown, not
distorted or sulcate when dry; urn to 4 mm, 4--5:1; operculum inclined-rostrate, about 1/5\x
length of urn; exostome to 500 \um,
teeth long and very narrow with base only 50 \um wide, evenly contracted from
base without a shoulder; outer surface of exostome horizontally to obliquely
striate basally but becoming smooth to lightly papillose distally, pale yellow,
erect when dry; endostome smooth with
segments shorter than the exostome, very narrow but perforate along mid-line,
arising from a short basal membrane with cilia paired and appendiculate. Spores
smooth, 10--14 \um. Forming large colonies over rock walls and boulders
especially along streams, coastal mountains influenced by summer fog,
infrequent on basal burls of broadleaf trees, especially (Umbellularia); moderate elevations; Bestia longipes is
usually recognized by the pattern of erect to decumbent, frondose
branching. The frondose branches have
an elongate main axis with markedly shorter branch axes so there may be a
10:1 ratio of length to width. Bestia longipes is infrequently
collected with sporophytes, which may be difficult to see at first. This is due to the fact that sporophytes
initiate from perichaetia inserted on the ventral side of the decumbent main
axis thereby seta elongation parallels the stem and substrate as opposed to being
elevated. Additionally the sporophytes are not much longer than the closely
associated ultimate branch axes. Bestia
can be confused in the field only with Isothecium,
especially hydrated plants when both occur mixed. Bestia can be readily distinguished by serrations only distally
in the leaf, cells uniform throughout, generally 4:1, and costa prominent, extending
to within 10 cells of the apex. In Isothecium,
the serrations occur over a much larger portion of the leaf margin, the alar
region is distinctly lighter colored compared to the remainder of the lamina,
and juxtacostal cells approach 8:1. 2. ISOTHECIUM Brid., Bryol.
Univ. 2: 255. 1829 * [Greek isos, equal, and theke, capsule, alluding to symmentric
capsule] W.
B. Schofield† Plants
slender, in often dense, pendent wefts. Stems creeping, irregularly branched,
bearing distant broadly triangular, 1-costate or ecostate weakly toothed
decurrent leaves and blunt foliose to deltoid pseudoparaphyllia; secondary
branches often dendroid and regularly or irregularly branched, these
sometimes producing flagelliferous terminal shoots with reduced leaves. Primary stem leaves similar to those
of the branchlets, erect-spreading to strongly imbricate, occasionally
wide-spreading, dark green to brownish green, occasionally yellow-green to
bronze, somewhat glossy. Branchlet
leaves ovate to ovate-lanceolate, coarsely toothed, especially in apex,
margins plane to slightly undulate; costa single, often ending in a spine
abaxially; leaf cells smooth or prorulose abaxially, usually hexagonal to
elongate in most of lamina; alar cells in a well-defined, sometimes excavate
group of smaller shorter cells, varying in number, often grading into
adjacent elongate cells. Seta elongate, red-brown, smooth,
1--3 cm. Capsule cylindric, rarely
erect, mostly suberect or inclined, 1.5--3 mm; operculum conic; peristome
double, yellow. Calyptra
naked or sparsely hairy. Species 11 (5 in the flora): Northern Hemisphere in
mainly temperate to subtropical climates, Atlantic and Pacific North America,
Europe, SELECTED REFERENCES: Allen, B. H. 1982. Isothecium alopecuroides new to 1.
Secondary stem system irregularly branched, strongly julaceous when dry;
leaves with extensive triangular region of alar cells extending to 1/3\x leaf
length . . . 3. Isothecium cristatum 1.
Secondary stem system irregularly or regularly branched, not julaceous, or if
julaceous regularly branched, sometimes dendroid; leaves with reduced
(sometimes excavate) region of alar cells extending less than 1/10\x leaf
length. 2.
Plants robust, 3--15 cm, secondary system often markedly dendroid and with
pronounced stipe, rarely producing tertiary flagelliferous branches,
irregularly branched . . . 2. Isothecium cardotii 2.
Plants medium-sized, usually 3--9 cm, less differentiated, often producing
attenuate tertiary branches, sometimes closely pinnate-branched. 3.
Secondary systems pinnate, julaceous, glossy . . . 5. Isothecium
stoloniferum (in part) 3.
Secondary systems pinnate or irregularly branched, not strongly glossy . 4.
Secondary systems often pinnate, often producing extensive flagelliform
branches . . . 5. 4.
Secondary systems irregularly branched, not producing flagelliform branches. 5.
Branch leaves of secondary system blunt, with few teeth, shoots julaceous . .
. 1. Isothecium alopecuroides 5.
Branch leaves of secondary system long-acuminate, sharply toothed, julaceous
or not. 6.
Branches of secondary system blunt, but leaves acuminate; julaceous . . . 5. Isothecium
stoloniferum (in part) 6.
Branches of secondary system sharp-tipped, leaves acuminate and
erect-spreading or wide-spreading, not julaceous . . . .4. Isothecium myosuroides 1. Isothecium alopecuroides (Lamark ex Dubois) Isoviita, Ann. Bot. Fenn. 18: 202. 1981 Hypnum
alopecuroides Lamarck ex Dubois, Méth. Éprouv.,
228. 1803; Isothecium myurum Bridel ex Amann Plants
robust, yellowish to brownish green, with secondary stems
somewhat pendent, irregularly branched, 1--5 cm, stipe short or not evident, the branches of equal length (1--2 cm). Stems
irregularly branched. Primary stem
leaves distant, small, broadly to narrowly triangular, squarrose, with
attenuate and sometimes secund apex; costa short or ecostate; margins weakly
or not toothed; alar cells weakly differentiated; pseudoparaphyllia scarce,
foliose, pointed or blunt. Branchlet
leaves of secondary system ovate to ovate-lanceolate, 2--3 cm, imbricate
to subjulaceous when dry; cucullate, costate to 2/3\x leaf length, tapering
abruptly to blunt, slightly toothed apex, otherwise with entire margins;
cells smooth, 7--12:1 in most of lamina, shorter and rhomboid at apex; alar
cells pigmented, rounded to 4--6-sided, in an often excavate group confined
to base just distal to marginal insertion.
Seta 0.8--1.2 cm. Capsule erect, 2--2.4 mm. Rock;
moderate elevations; Nfld.; Ont.; The
description of Isothecium alopecuroides
is based on European specimens. It
has been reported from 2.
Isothecium cardotii
Kindberg in Macoun & Kindberg, Bull. Torrey Bot. Club
17: 278. 1890 E F Plants
robust, dark green or yellow to golden green, secondary
system often forming extensive slightly glossy mats, 3--15 cm. Stems creeping, irregularly branched;
secondary stems rigid or soft, red-brown, usually stipitate, the stipe
bearing distant leaves resembling those of primary systems, producing a
frond-like pinnate branch-system (sometimes plumose), or irregularly
branched, resulting in a somewhat dendroid system, arching downward when dry,
sometimes flat dendroid and erect from substratum when humid, lateral
branches not flagelliform; pseudoparaphyllia scarce, broadly foliose, blunt
or with a few short teeth. Primary
stem leaves small, broadly to narrowly triangular, often of colorless,
distant squarrose leaves; margins entire to weakly toothed apex extending
abruptly as an attenuate point, sometimes falcate; costa present or absent;
alar region small, slightly decurrent with smaller, mainly truncate
rectangular cells, extending into decurrency. Branchlet leaves ovate-lanceolate, tapering to long (sometimes
twisted) acumen, margins coarsely toothed distally, basal margins nearly
entire to weakly toothed; costa broad at base tapering toward apex,
occasionally weakly forked, sometimes ending in an abaxial spine, extending
to 2/3\x leaf length; alar cells in well-defined, sometimes excavate group of
truncate, shorter, rectangular cells, extending ca. 1/3 distance across base
but less than 1/10\x leaf length. Seta 1--2 cm. Capsule inclined, 1.5--2 mm.
Tree
trunks, rocky cliffs and boulders; low and moderate elevations; B.C.; Isothecium
cardotii is clearly related to I.
stoloniferum and can grow mixed with it, but is readily separated in size
alone in most specimens, although occasionally specimens of I. stoloniferum are of similar size
and this creates difficulties, but the presence of flagelliform branches 3.
Isothecium cristatum
(Hampe) H. Robinson, Bryologist 65: 95. 1962
E Leptohymenium
cristatum Hampe, Linnaea 30: 459. 1860;
Isothecium aggregatum (Mitten)
A. Jaeger; I. brewerianum (Kindberg)
Kindberg; I. howei Kindberg; Plants
medium-sized, dark green to brownish green. Stems irregularly branched, 1--5 cm,
glossy, julaceous when dry, nearly of equal lengths with slight suggestion of
a differentiated main stem except for a short stipe; branches curved downward
when dry, attenuate at tips, with no tendency to produce flagelliform
branches; pseudoparaphyllia foliose, deltoid, pointed. Primary stem leaves squarrose, broadly ovate, tapering abruptly
to attenuate point (sometimes falcate); margins weakly toothed to nearly
entire; costa weak or obscure and confined to leaf base; alar cells in
well-differentiated triangular group to 1/4\x leaf length. Brachlet leaves ovate to
ovate-lanceolate, 1.5--2 mm, plane, sometimes tapering abruptly to
coarse-toothed apex; basal margins entire; costa slender; alar cells
variable, usually a triangular area of shorter cells to 1/4\x leaf length,
larger leaves of lateral branches gradually tapered and with more pronounced alar
cells. Seta 1--2 cm. Capsule
erect to suberect, 1--2 mm. On
trees and rocks, or logs, mainly in summer-dry-winter wet climates. Endemic
to western North America, from Isothecium
cristatum is a distinctive species with strongly julaceous shoots.
The branch leaves of the secondary stems show a triangular area of alar cells
most distinctly, which separates it from other species of the genus. From Pterogonicum
gracile, this feature as well as the single costa is immediately distinguishing.
Antheridial plants have small bulbiform perigonia on main stem as well as
lateral branches, and perichaetia are generally on main shoots 4.
Isothecium myosuroides
Bridel,
Bryol. Univ. 2: 369. 1827 Pseudisothecium
myosuroides (Bridel) Grout Plants
yellowish to dark green, with creeping primary system,
radiculose. Stems irregularly
branched; secondary system forming soft mats of often proliferating,
sometimes stipitate, arching shoots, 2--4 cm, stipes short to absent;
branching irregular, sometimes close, with branch-system arching downward,
frequently producing attenuate 2--5 cm flagelliform branches, with narrower
smaller leaves; pseudoparaphyllia foliose, sometimes toothed, often blunt and
broad, infrequent. Primary stem leaves
distant, broadly or narrowly triangular, squarrose, sometimes secund with
attenuate points, nearly entire; costa present or absent; alar cells barely
differentiated. Branchlet leaves ovate to ovate-lanceolate,
1--1.5 mm; margins plane, toothed throughout, most strongly in attenuate
apiculus; costa to 2/3\x leaf length; alar cells in well-defined small group
at margin of leaf insertion, sometimes excavate, mainly of shorter cells than
lamina, except in shoot decurrent portion.
Seta 1--2 cm. Capsule 1--1.5 mm. Predominately
rock, cliffs, boulders usually in shaded sites in forests, also occasionally
epiphytic and at tree bases, mostly low to moderate elevations; N.S., Nfld., P.E.I., The
eastern representation of Isothecium
myosuroides is clearly the same as the European species, but the relationship
of western North American material is unclear. K. Ryall et al. (2005) supported this
conclusion with molecular data, but the situation concerning some 5.
Isothecium stoloniferum
Bridel,
Bryol. Univ. 2: 371. 1827 E Isothecium
acuticuspis (Mitten) Macoun
& Kindberg, I. brachycladon
Kindberg, I. myurellum
Kindberg, Plants
dull to extremely glossy, brownish green, dark green, pale green to
golden-green, occasionally bronze-brown, radiculose, with rhizoids in tufts. Stems irregularly branched; secondary
branch system often acutely pinnate, elongate, 2--9 cm, with short
stipe-bearing costate leaves smaller and more abruptly tapering to the
toothed apex than those of tertiary branches, sometimes irregularly branched,
these often producing flagelliform tips to 5 cm and with smaller, more
distant leaves with alar cells weakly differentiated; pseudoparaphyllia
foliose, blunt or deltoid, blunt or toothed. Primary stem leaves variable in orientation, generally
erect-spreading, but occasionally widely divergent or strongly imbricate,
distant, broadly triangular, squarrose with attenuate tips and nearly entire
margins; costa present or absent; alar cells in small group at base, not
decurrent, isodiametric except in decurrent
portion, where elongate. Branchlet leaves lanceolate to
ovate-lanceolate or ovate, usually long-attenuate to a narrow acumen; margins
coarsely toothed throughout; costa single, often ending in a spine; leaf
cells smooth (sometimes prorulose abaxially), elongate; alar region confined
to base near margin just distal to insertion, often excavate, of mainly isodiametric cells, except those at insertion where
rectangular. Seta 1--1.5 cm. Capsule 1.5--2 mm. Epiphytic
on tree trunks and branches, occasionally on shrubs, exposed or shaded rock
cliffs, boulder slopes, forests or open sites; low to moderate elevations (0--1500
m); Alta., B.C.; Isothecium
stoloniferum is an extremely variable species
strongly related to the less variable A
third morphotype is the predominant one of In
considering the I. stoloniferum complex,
it is strongly suggested that 3. TRIPTEROCLADIUM A. Jaeger, Ber.
Thätik. St. Gallischen Naturwiss. Ges. 1877--78: 484.
1880 * [Greek tri, three; pteron, wing; klados, branch;
alluding to perceived tripinnate branching] Howard
A. Crum† Plants
rather small, shiny. Stems
irregularly 1(--2)-pinnate, the branches of unequal length; pseudoparaphyllia
broadly rounded-foliose. Stem leaves
oblong-ovate, broadly acuminate, decurrent; margins
plane or somewhat recurved near the base, entire or ± serrulate; costa short
and double to single and sometimes extending nearly to mid leaf; distal cells
linear, thick-walled, smooth; alar cells small and subquadrate
in rather sizable groups. Branch
leaves smaller, more broadly acuminate, irregularly
serrate distally. Perichaetial leaves
pale, sheathing at base, abruptly short-subulate,
entire or nearly so. Seta red,
smooth. Capsule suberect and ±
curved to horizontal; annulus well differentiated; operculum conic; stomata
in the short neck; exostome teeth pale yellow, cross-striolate below;
endostome a high basal membrane with narrow, densely papillose segments
perforate along the keel, cilia single, nodose. Calyptra not seen. Species 1 (1 in the flora): w Tripterocladium
is a unispecific genus in the past moved in and out of the
Lembophyllaceae, but molecular data confirm its position in the family as
closely related to Bestia. SELECTED REFERENCES Crum, H. A. 1987. Bestia, Tripterocladium, and Isothecium:
an explication of relationships. Bryologist 90: 40--42. 1.
Tripterocladium leucocladulum (Müller Hal.) A.
Jaeger, Ber. St. Gallischen
Naturwiss. Ges. 1877--78:
484. 1880 E F Hypnum
leucocladulum Müller Hal., Flora
58: 79. 1875 Plants
slender, pale yellow, green, or brownish. Stems erect-ascending, often filiform. Leaves of stem imbricate when dry,
erect-spreading when moist, 1.1--1.2 mm; costa short and double or single and
extending nearly to mid leaf; distal cells linear, thick-walled, smooth; alar
cells small and subquadrate in rather sizable
groups; leaves of branches 0.4--0.8 mm. Seta
1--1.5 cm. Capsules 1.5--2 mm. Spores papillose, 12--19 \um. Dry, shaded cliffs and boulders; B. C.; |