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BFNA Title: Mielichhoferiaceae |
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Illustrations,
click here. XX. Mielichhoferiaceae
Schimper A. Plants acrocarpous, with sporophytes terminal
though sometimes appearing lateral because of subfloral innovations, or cladocarpous
with sporophytes lateral on short branches; tiny to robust, as scattered
individuals or forming small to large colonies, green, reddish, whitish, or
golden, often glossy when dry. Stems
short to long, 0.3--10 cm, sometimes somewhat complanate, unbranched or
irregularly forking when sterile, often bearing subfloral innovations when
with gametangia; rhizoids sparse to abundant, variously colored, smooth to
papillose, often bearing multicellular tubers. Leaves imbricate to slightly twisted when dry, erect to
erect-spreading when wet, broadly lanceolate, ovate-lanceolate, elliptical,
obovate, 0.3--2.5 cm, decurrent or not; margins plane or revolute, finely
serrate to serrulate near the apex, rarely entire, apex bluntly acute to
acute or shortly acuminate, costa ending well before the apex, subpercurrent
or percurrent, rarely shortly excurrent, transverse section with stereid band
single, usually well developed, occasionally reduced, generally with guide
cells; distal medial laminal cells elongate-hexagonal, rhombic, or
linear-rhomboidal, sometimes narrowly vermicular, with thin to thickened
walls, proximal cells usually short- or long-rectangular, thin-walled,
marginal cells undifferentiated, slightly narrower and longer than median
cells [sometimes abruptly differentiated as a distinct border, rarely red]. Specialized asexual reproduction
common, variously spherical to ovoid rhizoidal tubers or axillary filiform to
buliform gemmae. Sexual condition
dioicous, paroicous, autoicous, rarely synoicous, rarely variable within
species (polyoicous); perigonia and perichaetia terminal or lateral,
perichaetial leaves the same size as vegetative leaves, sometimes strongly
differentiated, narrowly to linear-lanceolate, or smaller, with weaker costa,
perigonia bud-like, perigonial leaves broadly ovate, concave and often brown
to reddish basally, acute to long-acuminate distally, acumen erect to
wide-spreading. Seta usually
1(2--5), variously colored, long, straight, bent to various degrees producing
inclined to pendulous capsules. Capsule
erect, or inclined to 180° from vertical, long-exserted, 1--5(--10) mm,
narrowly cylindrical to broadly pyriform, neck well differentiated, sometimes
as long or longer than the urn, stomata abundant in the neck, superficial to
immersed, exothecial cells near mouth quadrate or short-rectangular,
thick-walled, often reddish in 1--3 or more rows, median cells longer, short-
to long-rectangular with straight or sinuose walls, rarely isodiametric,
sometimes collenchymatous; annulus present, revoluble, or absent; operculum
convex, short to tall-conic, sometimes shortly rostrate; peristome
diplolepideous-alternate, rarely reduced to one layer or absent; peristomial
formula 4:2:4--6, sometimes 4:2:8; exostome white, pale yellow to brown, rarely
red, teeth triangular to lanceolate, rarely reduced or absent, trabeculate or
not, pitted or rarely papillose below, coarsely or rarely finely papillose
distally, endostome hyaline to yellow, segments narrow to wide, strongly to
weakly or scarcely keeled, rarely absent or reduced to a rudimentary
membrane, broadly to narrowly perforate or less commonly entire, basal
membrane low to high, sometimes scarcely exceeding the capsule rim, cilia
present or absent, 1--3, as long as the segments or shorter, nodulose or not.
Calyptra fugacious, cucullate,
smooth. Spores shed singly, 11--45
\um, smooth to papillose, pale yellow, tan or nearly hyaline, rarely darker. Genera 4,
species ca. 150 (3 genera, 36 species in the flora): worldwide, most speciose
in the Northern Hemisphere. The family
Mielichhoferiaceae includes species that were traditionally classified in the
Bryaceae, but are phylogenetically closer to the Mniaceae. In addition to the three North American
genera, the family probably includes Pseudopohlia
of Among Epipterygium species, the North
American E. tozeri is most similar
to species of Pohlia, especially
those of the Mniobryum group (the
latter sometimes treated as a subgenus of Pohlia
or as a separate genus). Tropical species of Epipterygium, including those of the Caribbean region and Mielichhoferia is especially diverse in South America
and is highly variable in peristome development ranging from double with
well-developed exostome and endostome, single and endostomial, double but
with the exostome short and barely exceeding the capsule rim, or double with
the endostome reduced to an inconspicuous membrane adherent to the exostome
teeth (two North American species). Mielichhoferia species are
gametophytically similar to small species of Pohlia, though they can be distinguished by a characteristic
whitish color and very small size. The main features distinguishing the two
genera are gametangia borne on short lateral shoots in Mielichhoferia and terminal in Pohlia. The perichaetial
leaves are as long or longer than vegetative leaves in Pohlia but are smaller, with more lax cells and shorter costa in Mielichhoferia. Mielichhoferia and Pohlia
grow in similar habitats. The three
genera of Mielichhoferiaceae were almost universally classified in the
Bryaceae until C. J. Cox and T. A. J. Hedderson (2003) published their ground-breaking
phylogenetic study in which they showed that these genera are more closely
related to the Mniaceae than Bryaceae. Since that time, their conclusion has
been repeatedly corroborated by additional data from all three genomic
compartments; nuclear, chloroplast, and mitochondrial. Phylogenetic resolution of precise
relationships between genera in the Mielichhoferiaceae versus Mniaceace in
the strict sense has, however, been more recalcitrant (C. J. Cox, pers.
comm.). Molecular evidence from multiple loci suggests that the
Mielichhoferiaceae may form early diverging lineages within the broader
Mniaceae clade and the family may therefore be paraphyletic. However, relationships are poorly supported
and reciprocal monophylly for Mielichhoferiaceae and Mniaceae cannot
presently be rejected. Very short branch lengths separating early diverging
lineages of Mielichhoferiaceae and Mniaceae suggest that a rapid radiation
early in the history of this clade may be the explanation for poor resolution
of phylogenetic relationships among the lineages. SELECTED
REFERENCES: Cox, C. J. and T. A. J. Hedderson. 2003. Phylogenetic
relationships within the moss family Bryaceae based on chloroplast DNA
evidence. J. Bryol. 25: 31--40. 1. Peristome appearing single and exostomial;
exostome teeth long; endostome rudimentary and inconspicuous, adherent to the
inside base of exostome teeth; perichaetia and perigonia on short lateral
branches .................................................................. 1. Mielichhoferia,
p. XX 1. Peristome clearly double; endostome with
well-developed segments from a low to high basal membrane or, if reduced,
with at least a basal membrane free from the exostome teeth; perichaetia
terminal, although sometimes appearing lateral because of subfloral
innovations. 2. Plants not complanate, without differentiation
of dorsal and lateral leaves; leaves not bordered by linear cells although
marginal cells sometimes slightly narrower than median cells, lanceolate to
ovate-lanceolate .................................. 2. Pohlia,
p. XX 2. Plants slightly complanate, with 1--3 rows
of dorsal leaves smaller than the lateral leaves; leaves bordered by 1--4
rows of ± differentiated linear, marginal cells, elliptic 3. Epipterygium, p. XX 1. Mielichhoferia
Nees, Bryol. Germ. 2(2): 179. 1831 * [For Mathias Mielichhofer, 1772--1847,
collector of the generitype specimen in the Salzburg Alps] Plants small to medium size, usually forming
± compact turfs. Stems short to long, 0.1--3 cm, unbranched or irregularly forking
when sterile, bearing lateral gametangia when fertile. Leaves erect to erect-spreading, unaltered on drying, narrowly
lanceolate to ovate-lanceolate, 0.4--1.5 mm, not decurrent; margins of
vegetative leaves plane [rarely recurved], finely serrate to serrulate near
the apex, rarely entire, acute or shortly acuminate, costa ending well before
the apex to subpercurrent; distal medial laminal cells elongate-hexagonal,
rhombic, or long-rhomboidal, with thin to thickened walls, proximal cells
usually short- or long-rectangular, thin-walled, gradually differentiated,
marginal cells undifferentiated. Specialized
asexual reproduction rare, occasionally of bulbiform gemmae. Sexual condition dioicous, rarely
autoicous or synoicous; perigonia and perichaetia on short lateral branches; perichaetial
leaves typically smaller than vegetative leaves, with laxer cells, lanceolate
to narrowly linear-lanceolate, with plane to slightly recurved margins;
perigonia budlike, perigonial leaves broadly ovate, concave, brown to reddish
proximally, shortly acute distally, acumen erect to erect-spreading. Seta single, variously colored, long,
straight or bent to various degrees producing slightly inclined capsules. Capsule erect or inclined to ca. 30°
from vertical, 1--4 mm, narrowly cylindrical to pyriform, neck
well-differentiated, shorter than the urn, stomata abundant in the neck,
superficial, exothecial cells near mouth quadrate or short-rectangular,
thick-walled, often reddish, in 1--3 rows, medial cells longer, short- to
long-rectangular with straight or slightly sinuose walls; annulus present and
revoluble; operculum convex, short to tall-conic; peristome single or double,
diplolepideous-alternate when both exostome and endostome present [rarely
absent]; peristomial formula typically 4:2:4; exostome, when present, white
to hyaline, teeth narrowly triangular to lanceolate, sometimes irregular,
often reduced or absent, weakly or not trabeculate, papillose to nearly
smooth, endostome hyaline, segments narrow, weakly or scarcely keeled,
sometimes absent, broadly to narrowly perforate or entire, basal membrane low
to high, sometimes scarcely exceeding the capsule rim, absent. Mielichhoferia is characterized by perichaetia and
perigonia borne on short lateral branches, perichaetial leaves that are often
smaller and with cells that are more lax relative to those of the vegetative
leaves, and frequent peristome reduction. The leafy stems themselves are much
like those of a small Pohlia. Species ca. 45
(3 in the flora); North America, South America, Europe, Africa, Selected references.
Shaw, A. J. and H. A. Crum. 1984. Peristome homology in Mielichhoferia with a taxonomic account of the North American
species. J. Hattori Bot. Lab. 57:
363--381. Shaw, A. J. and P. E. Rooks.
1994. Systematics of
Mielichhoferia Nees & Hornsch. (Bryaceae: Musci) I. Morphological and genetic analyses of M. elongata and M. mielichhoferiana.
Bryologist 97: 1--12. Shaw, A.
J. 1994. Systematics of Mielichhoferia (Bryaceae: Musci) II. Morphological variation among disjunct
populations of M. elongata and M. mielichhoferiana. Bryologist 97: 47--55. 1. Plants synoicous; peristome single; exostome
absent, endostome of narrow, weakly keeled, smooth segments ................................................................... 3.
Mielichhoferia shevockii 1. Plant dioicous; peristome double; exostome
of 16 papillose teeth, endostome of a rudimentary membrane adherent to the
inside of the exostome teeth. 2. Leaves dull dark green; distal
medial leaf cells with firm to thickened walls ....................... 2. Leaves pale whitish green; distal
medial leaf cells lax and thin-walled ............................... 1. Mielichhoferia mielichhoferiana
(Funck) Loeske, Stud. Vergleich. Morph. Phylog. Syst. Laubm., 126. 1910 Weissia mielichhoferiana Funck, Crypt. Gew. Fichtelgeb. 24: 2.
1817 [1818] Plants medium sized to robust, dull green. Leaves 0.6--1.3 mm, erect to ±
spreading or commonly slightly secund, lanceolate; distal medial cells
hexagonal to rhomboidal, 50--90 \um, moderately thick-walled. Specialized asexual reproduction
absent or rarely present as axillary bulbiform gemmae. Sexual condition dioicous. Capsule
erect or inclined to 20° from vertical; exostome teeth hyaline to whitish,
narrowly acute- or bluntly triangular, papillose; endostome hyaline, segments
absent, basal membrane rudimentary and adherent to the exostome teeth. Spores 15--22 \um, roughened. Capsules
mature summer (Jun--Aug). Rock crevices, soil banks, roadsides; low to high
elevations; Greenland; B.C., Calif., Colo.; Eurasia. Mielichhoferia
mielichhoferiana
is distinguished from M. elongata
by darker green, more elongate-lanceolate, dull leaves that are commonly
somewhat secund. The leaf cells of M. mielichoferiana are smaller and
narrower, hexagonal to rhombic, with thickened walls. The two species
hybridize when they grow in mixed colonies but are usually readily
distinguishable. Recombinant plants
that have genetic markers from both M.
mielichhoferiana and M. elongata,
presumably derived from hybrid sporophytes, have smaller, shorter, straighter
leaves (not secund) than typical M.
mielichhoferiana. They tend to have the dull, dark color of M. mielichhoferiana rather than the
pale whitish aspect of M. elongata.
Mielichhoferia mielichhoferiana is
much less common than M. elongata
in North America, and this is apparently so in 2. Mielichhoferia elongata (Hoppe & Hornschuch) Nees &
Hornschuch, Bryologia Germanica 2(2): 186. 1831 Weissia elongata Hoppe & Hornschuch, Musci Exotici
2: 102. 1819 Plants slender to medium size, whitish
pale-green. Leaves 0.5--1.1 mm,
erect to ± spreading, lanceolate; distal medial cells rhomboidal, 50--100
\um, thin-walled. Specialized asexual
reproduction absent. Sexual
condition dioicous. Capsule
erect or inclined to 20° from vertical, [peristome sometimes rudimentary]
exostome teeth hyaline to whitish, narrowly acute- or bluntly triangular,
papillose; endostome hyaline, segments absent, basal membrane rudimentary and
adherent to the exostome teeth. Spores
15--22 \um, roughened. Capsules mature spring (Apr--Jun).
Rocks and soil, often on substrates naturally enriched with heavy metals;
mine tailings; low to high elevations; Greenland; Alta., B.C., Nfld. and
Labr., Nunavut; Alaska, Calif., Colo., Maine, Mich., Mont., N.Y., N.C.,
Tenn., Europe; Asia. Mielichhoferia elongata grows in dense, compact turfs with a
whitish or sometimes bluish color that makes this species easily
recognizable. The leaf cells are thin-walled, giving the plants a softer
texture than M. mielichhoferiana.
The species might be mistaken for a Pohlia,
but the turfs are usually more compact and the plants are smaller; when
fertile the gametangial buds are lateral and the sporophytes are erect or
nearly so. There is substantial variation in development of the exostome
teeth but North American plants have relatively long, rather coarsely
papillose segments. This species
is one of the so-called copper mosses because it often grows on mineral
enriched substrates, although not always with copper. Like other copper
mosses, M. elongata has a broad and
highly disjunctive geographic range but is nowhere common. In North America, it is most frequent in
the Rocky Mountains of Colorado, where it occurs on disused mine tailings,
especially around the towns of Ouray and Silverton in 3. Mielichhoferia shevockii (A. J. Shaw) A. J. Shaw, Bryologist XXX(YY): . Schizymenium shevockii A. J. Shaw, Syst. Bot. 25: 190, figs.
1--9. 2000 Plants medium sized, dull green. Leaves 0.8--1.3 mm, erect to ±
spreading, lanceolate; distal medial cells hexagonal to rhomboidal, 60--110
\um, moderately thick-walled. Specialized
asexual reproduction absent. Sexual
condition dioicous. Capsule
erect or inclined to 30° from vertical; exostome absent; endostome hyaline to
pale yellow, segments short, irregular, narrow, slightly keeled, not
perforate, basal membrane scarcely exceeding the capsule rim. Spores 15--20 \um, finely roughened. Capsules
mature spring (Apr--Jun). Known from only three proximate sites, soil and
rock; The
gametophytes of M. shevockii look
much like those of genetically recombinant intermediates between M. elongata and M. mielichhoferiana derived from hybrid sporophytes, but can be
readily distinguished by DNA markers.
The gametophytes of S. shevockii,
like those intermediate between M.
elongata and M. mielichhoferiana,
have dull leaves that are a little more pale than is typical of pure M. mielichhoferiana, and are
relatively small like those of M.
elongata. Unlike those two species, M.
shevockii is synoicous and has a single endostomial peristome. The segments are nearly smooth. In the original description of M. shevockii (as Schizymenium), the sexuality was incorrectly described as
dioicous. |
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2. Pohlia
Hedwig, Sp. Musc. Frond., 171. 1801 * [For Johann E. Pohl, 1782--1834,
physician of Plants tiny to robust, occurring as scattered
plants or as deep and sometimes extensive turfs. Stems short to long, 0.1--10 cm, unbranched or irregularly
forking when sterile, often bearing subfloral innovations when with
gametangia. Leaves erect to
erect-spreading, rarely wide-spreading when wet, unaltered or sometimes
slightly contorted on drying, narrowly lanceolate to ovate-lanceolate [rarely
linear-lanceolate], 0.4--3 mm, decurrent or not; margins of vegetative leaves
plane or rarely revolute, finely serrate to serrulate near the apex, rarely
entire, acute or shortly acuminate, costa ending well before the apex,
subpercurrent or percurrent, rarely shortly excurrent; distal medial laminal
cells elongate-hexagonal, rhombic, or linear-rhomboidal, sometimes narrowly
vermicular, with thin to thickened walls, proximal cells usually short- or
long-rectangular, thin-walled, gradually differentiated, marginal cells
undifferentiated or slightly narrower and longer than median cells. Specialized asexual reproduction
common, variously as spherical to ovoid rhizoidal tubers or axillary filiform
to bulbiform gemmae. Sexual condition
dioicous, paroicous, rarely autoicous or synoicous, rarely variable within
species (polyoicous); perigonia and perichaetia terminal, rarely the
perigonia lateral (P. longibracteata);
perichaetial leaves the same size as vegetative leaves or sometimes strongly
differentiated, lanceolate to narrowly to linear-lanceolate, with recurved
margins; perigonia bud-like, perigonial leaves broadly ovate, concave and
often brown to reddish proximally, acute to long-acuminate distally, acumen
erect to wide-spreading. Seta
1(2--5), variously colored, long, straight, bent to various degrees producing
inclined to pendulous capsules. Capsule
erect or inclined to 180° from vertical, 1--6[--10] mm, narrowly cylindrical
to broadly pyriform or urceolate, neck well differentiated, sometimes as long
or longer than the urn, stomata abundant in the neck, superficial to
immersed, exothecial cells near mouth quadrate or short-rectangular,
thick-walled, often reddish in 1--3 or more rows, medial cells longer, short-
to long-rectangular with straight or sinuose walls; annulus present and
revoluble or absent; operculum convex, short to tall-conic, sometimes shortly
rostrate; peristome diplolepideous-alternate [rarely absent]; peristomial
formula 4:2:4--6, rarely 4:2:8; exostome white, pale yellow to brown,
sometimes dark reddish brown, teeth triangular to lanceolate, sometimes
irregular [rarely reduced or absent], trabeculate or not, pitted or rarely
papillose basally, coarsely or rarely finely papillose distally, endostome
hyaline to yellow, segments narrow to wide, strongly to weakly or scarcely
keeled, rarely reduced to rudimentary projections, broadly to narrowly
perforate or less commonly entire, basal membrane low to high, sometimes
scarcely exceeding the capsule rim, cilia present or absent, 1--3, as long as
the segments or shorter, nodulose or not. Species ca. 85
(32 species in the flora): North America, Central America, South America,
Europe, Asia, Africa, SELECTED
REFERENCES. Ochi, H. 1959. A revision of the Bryaceae in 1. Plants with axillary gemmae. 2.. Gemmae with broadly laminate (leaf-like)
leaf primordia 3.. Gemmae in clusters of 4--many per
leaf axil, narrowly elongate to oblong-bulbiform or obconic 4.. Gemmae linear, extending beyond the
leaves ...................... 15.
Pohlia tundrae 4.. Gemmae oblong, obconic, or
spheroidal. 5. Gemmae spheroidal to oblong,
occasionally ovate ........ 20. Pohlia bulbifera 5. Gemmae obconic, distinctly narrowed to
the base ..... 19. Pohlia andalusica 3.. Gemmae 1--2 per leaf axil, bulbiform. 6.. Gemmae ovate; leaf primordia small,
stiffly triangular, mainly restricted to gemma apex or rarely 1--2 primordia
occurring more proximally .......... 17.
Pohlia filum 6.. Gemmae elongate-bulbiform; leaf
primordia larger, flexuose, at the gemma apex and also commonly more
proximally. 7. Plants dull, leaves slightly contorted
when dry ..... 16. Pohlia rabunbaldensis 7. Plants shiny, leaves unaltered when
dry. 8.. Leaves green or tinged with red,
loosely erect to erect-spreading .............. 8.. Leaves pale whitish, stiffly erect ........................... 18.
Pohlia beringiense 2.. Gemmae with peg-like leaf primordia. 9.. Plants shiny when dry. 10. Gemmae linear-vermicular, with 1--2
elongate, erect leaf promordia 10. Gemmae oblong, with 1--5 short, incurved
leaf primordia 24.
Pohlia andrewsii 9. Plants dull when dry. 11. Gemmae oblong, knobby in outline because of
angular external cells, without leaf primordia ....................................................................... 25. Pohlia flexuosa 11. Gemmae spheroidal, obconic, or linear, not
knobby in outline, external cells rounded, with leaf primordia 12.. Gemmae spheroidal ..................................... 22. Pohlia camptotrachela 12.. Gemmae oblong or obconic to linear,
clearly longer than wide. 13. Gemmae oblong or obconic, with 2--5
conspicuous leaf primordia 13.. Gemmae narrowly linear, with 1(--2)
inconspicuous leaf primordia 25. Pohlia flexuosa 1.. Plants lacking axillary gemmae. 14. Leaf margins recurved from base to apex. 15.. Leaves narrowly long-lanceolate,
median leaf cells long- to linear-hexagonal, with thin, non-porose walls .................................................... 8. Pohlia crudoides 15.. Leaves ovate-lanceolate,
median leaf cells rhombic, with thickened ± porose walls ......................................................................................... 10.
Pohlia cardotii 14. Leaf margins plane (perichaetial leaves with
± recurved margins). 16.. Plants
paroicous. 17.. Plants
polysetous .................................................. 6. Pohlia robertsonii 17.. Plants with one seta per
inflorescence. 18.. Plants with lax, hexagonal,
thin-walled median cells more than 10 \um wide ............................................................... 7. Pohlia obtusifolia 18.. Plants with firm, hexagonal to
rhomboidal, ± thick-walled cells less than 10 \um wide. 19.. Plants very shiny when dry, leaf
cells linear-vermicular. 20.. Leaves broadly lanceolate to
elliptic; endostome segments broadly perforate....................................... 5. Pohlia cruda 20.. Leaves long triangular-lanceolate;
endostome segments narrowly or not all perforate, cilia absent 3.
Pohlia longicollis 19.. Plants dull when dry, leaf cells
short- to long-hexagonal. 21.. Neck of capsule shorter than the urn;
endostome segments broadly perforate, cilia present.................. 1. Pohlia nutans 21.. Neck of capsule as long or longer
than the urn; endostome segments narrowly or not all perforate, cilia absent 2.
Pohlia elongata 16.. Plants
dioicous (very rarely autoicous). 22.. Capsules
erect or nearly so, endostome segments narrow, scarcely keeled, hardly
tapered, cilia absent .......................................... 11.
Pohlia erecta 22.. Capsules inclined 50--180° from
vertical, endostome segments broader, keeled, tapered apically, cilia present
or absent. 24.. Leaves long-decurrent, median leaf
cells leaf cells rhombic to rhomboidal, less than 10 \um wide ....................... 9. Pohlia
ludwigii 24.. Leaves not or scarcely decurrent,
median leaf cells various but if leaves decurrent, median cells more than 12
\um wide. 25.. Capsules narrowly cylindrical,
pendent 50--90° from vertical; endostome segments narrowly perforate, cilia
absent .................. 25.. Capsules pyriform to urceolate,
pendent 150--180° from vertical; endostome segments broadly perforate, cilia
present. 26.. Annulus present; exothecial cells
longer than wide. 27... Plants dull green, leaf cells
shortly hexagonal ................ 27... Plants pale or shiny green, leaf
cells long- to linear-rhomboidal. 28.. Plants very glossy green; leaves
coarsely serrate at apex; median leaf cells 110--180 \um 13.
Pohlia 28.. Plants pale green, not or slightly
glossy; leaves serrulate at apex; median leaf cells 65--120 \um ......................................... 12.
Pohlia lescuriana 26.. Annulus absent; exothecial cells
little or no longer than wide. 29... Median leaf cells laxly rhomboidal,
12 \um or more wide. 30.. Plants tiny (2--6 mm), reddish;
stomates superficial or slight immersed; cells of perichaetial bracts
narrower than those of vegetative leaves 30.
Pohlia atropurpurea 30.. Plants small to robust (5--80 mm);
yellow-green to whitish, rarely (when very robust) with some red pigmentation;
stomates deeply immersed; perichaetial leaf cells broad and lax, like those
of vegetative leaves 31... Plants pale whitish (sometimes
tinged with red when very robust), 1--8 cm 31. Plants deep green, 0.5--1 cm high ............... 29... Median leaf cells linear-hexagonal
to linear-rhomboidal, 6--10 \um wide. 32.. Plants medium size to robust; leaves
wide-spreading; perigonial bracts long-acuminate, to 6 mm.......................... 26. Pohlia
longibracteata 32.. Plants slender; leaves erect to
erect-spreading; perigonial bracts shorter, 1.5--3 mm. 33... Stomates deeply immersed. 34.. Stems cherry-red; leaves erect,
glossy-green .................... 31. Pohlia
vexans 34.. Stems green to pale orange or pink;
leaves spreading, dull, often reddish 33... Stomates superficial or slightly
immersed. 35. Stems elongate, to 1 cm; leaves erect,
whitish; median cells of vegetative leaves linear, 6--9 \um wide 35. Stems
short, 2--6 mm, loosely spreading (except perichaetial leaves); median leaf
cells of sterile plants and of proximal vegetative leaves broadly hexagonal,
10--12 \um wide |
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1.
Pohlia nutans (Hedwig)
H. Lindberg, Musci Scand., 18. 1879 Webera nutans
Hedwig, Sp. Musc. Frond.,
168. 1801; Pohlia sphagnicola (Bruch & Schimper) Lindberg
& Arnell Plants small to robust, dull green or rarely
reddish. Stems 0.5--3.5(--10) cm. Leaves to 2 mm, erect to ± spreading,
lanceolate to ovate-lanceolate, acute; margins subentire or more commonly
serrulate to serrate in the distal 1/3; costa subpercurrent, percurrent, or
rarely shortly excurrent, broad; distal medial cells hexagonal to rhomboidal,
50--90 \um, moderately thick-walled. Specialized
asexual reproduction absent. Sexual
condition paroicous, rarely dioicous; perichaetial leaves somewhat
differentiated, ± long-lanceolate, perigonial leaves ovate-lanceolate. Seta orange to orange-brown. Capsule inclined 80--100° from vertical,
orange to orange-brown, slenderly pyriform, neck ca. 1/2 the urn length; exothecial cells
elongate-rectangular, walls straight; stomata superficial; operculum conic;
exostome teeth yellow to orange-brown, acute-triangular, pitted basally,
coarsely papillose distally; endostome hyaline (to rarely orange-brown),
basal membrane 1/2\x the exostome length, segments broadly keeled, broadly
perforate, cilia short to long. Spores
16--22 \um, finely to distinctly roughened. Capsules mature spring (Apr--Jun).
A common species of soil banks, logs, tree bases and sometimes disturbed
places; low to high elevations; Greenland; Alta., B.C., Man., N.B., Nfld. and
Labr., N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Ala., Alaska,
Ariz., Ark., Calif., Colo., Conn., Del., D.C., Fla., Ga., Idaho, Ill, Ind.,
Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., Mo., Mont.,
Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Oreg., Pa.,
R.I., S.C., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.;
Mexico; Eurasia; Africa; Australia. Pohlia nutans is the most common species of the
genus in North America, as it is in The peristome
of P. nutans is as well developed
as any in Pohlia, with long
tapered, trabeculate exostome teeth and well-developed endostomes with
broadly keeled, widely perforate segments and short to long cilia. The leaf
cells are short-to elongate-hexagonal with thickened walls. The only other North American species with
relatively thick-walled leaf cells is P.
elongata. The cells of P. elongata are typically longer but
there is extensive overlap in sizes. Plants without sporophytes have
generally been named P. nutans
although some such collections, from montane regions, may actually be P. elongata. Sporophytes of P. elongata have longer necks, as long
as or longer than the urns, and reduced peristomes with endostome segments
narrowly split along the keel and (usually) no cilia. 2. Pohlia elongata
Hedwig, Sp. Musc. Frond., 171. 1801 Pohlia polymorpha Hoppe & Hornschuch ex Hornsuch; P. acuminata Hoppe & Hornschuch ex
Hornschuch; P. elongata var. greenii (Brid.) A.J. Shaw Plants small, dull green. Stems 0.5--2.5 cm. Leaves to 2 mm, erect to ± spreading,
lanceolate to ovate-lanceolate, acute; margins subentire or more commonly
serrulate to serrate in the distal 1/3; costa subpercurrent or percurrent;
distal medial cells hexagonal to rhomboidal, 45--85 \um, moderately thick-walled.
Specialized asexual reproduction
absent. Sexual condition
paroicous, autoicous, rarely dioicous; perichaetial leaves somewhat
differentiated, ± long-lanceolate; perigonial
leaves in dioicous plants shortly and broadly ovate. Seta orange to orange-brown. Capsule
inclined 10--90° from vertical, stramineous, orange, or orange-brown, shortly
to longly and slenderly pyriform, neck ca. 1/2--1\x the urn length;
exothecial cells elongate-rectangular, walls straight; stomata superficial;
operculum conic; exostome teeth yellow to brown, acute-triangular, pitted
basally, coarsely papillose distally; endostome hyaline, basal membrane
barely exceeding the capsule rim or up to 1/3\x the exostome length, segments
narrowly or not keeled, narrowly perforate to entire, cilia absent to
rudimentary, rarely long. Spores
16--23 \um, distinctly roughened. Capsules
mature spring to summer (Apr--Jul). Humus-rich soil banks, along streams and
paths, tree bases; moderate to high elevations; B.C., N.B., Nfld. and Labr.,
N.W.T., N.S., Ont., Que., Yukon; Ala., Alaska, Ariz., Calif., Colo., Ga.,
Iowa, Maine, Mich., Minn., Mont., N.H., N.Mex., N.Y., N.C., Ohio, Tenn.,
Utah, Vt., Va., Wash., Wis.; Mexico; Central America, South America; Eurasia;
Africa; Pacific Islands; Australia. Pohlia elongata is widespread in 3. Pohlia longicollis (Hedwig) H. Lindberg, Musci Scand., 18.
1879 Plants medium-size, shiny green to
yellow-green. Stems 0.8--3.5 cm. Leaves 1.5--2.6 mm, erect to ±
spreading, long-lanceolate, acute; margins serrulate to serrate in the distal
1/3; costa subpercurrent or percurrent; distal medial cells
linear-rhomboidal, 80--140 \um, vermicular, thin-walled. Specialized asexual reproduction absent. Sexual condition paroicous; perichaetial leaves gradually
differentiated, ± long-lanceolate. Seta
orange to orange-brown. Capsule
inclined 10--90° from vertical, stramineous to orange-brown, longly and
slenderly pyriform, neck ca. 1/2--1\x the urn length; exothecial cells
elongate-rectangular, walls straight; stomata superficial; operculum conic;
exostome teeth yellow to brown, acute-triangular, pitted basally, coarsely
papillose distally; endostome hyaline, basal membrane 1/4--1/2\x the exostome
length, segments narrowly keeled, narrowly perforate to entire, cilia absent
to rudimentary. Spores 16--23 \um,
distinctly roughened. Capsules
mature spring to summer (Apr--Jul). Humus-rich soil banks, along streams and
paths; moderate to high elevations; Greenland; Alta., B.C., Nfld. and Labr.,
N.W.T., Ont., Yukon; Alaska, Mont., N.H., N.Y., N.C., Oreg., Pa., Tenn.,
Utah, Va., Wyo.; Mexico; Eurasia. Pohlia longicollis is a handsome species characterized by
relatively large, very shiny, long-lanceolate leaves. The plants are bisexual (paroicous) and
commonly form sporophytes although P.
longicollis is relatively rare in 4. Pohlia
bolanderi (Lesqereux) Brotherus in A. Engler & K. Prantl,
Nat. Pfl. 1(3): 548. 1903 Bryum bolanderi Sullivant, Mem. Plants small to medium-size, somewhat shiny
green or sometimes pale whitish; Stems
0.3--1.5 cm. Leaves 0.8--1.6 mm,
erect to ± spreading, lanceolate to narrowly ovate-lanceolate, acute; margins
serrulate to serrate in the distal 1/3; costa subpercurrent or percurrent;
distal medial cells linear-rhomboidal, 55--100 \um, thin-walled. Specialized asexual reproduction
absent. Sexual condition dioicous;
perichaetial leaves slightly differentiated; perigonial leaves ovate,
short-acuminate. Seta
straw-colored to orange-brown. Capsule
inclined 35--90° from vertical, straw-colored to orange-brown, slenderly
pyriform, neck ca. 1/2--1\x the urn length; exothecial cells
elongate-rectangular, walls straight; stomata superficial; operculum conic;
exostome teeth yellow to yellow-brown, acute-triangular, pitted basally,
coarsely papillose distally; endostome hyaline, basal membrane 1/4--1/2\x the
exostome length, segments narrowly keeled, narrowly perforate to entire,
cilia absent to rudimentary. Spores
16--26 \um, distinctly roughened. Capsules mature summer (Jun--Aug).
Rather dry alpine soil or soil-filled rock crevices; low to high elevations;
B.C.; Alaska, Calif., Colo., Nev., N.Mex., Oreg., Utah, Wash., Wyo.; Europe. Pohlia bolanderi has glossy to whitish leaves. Glossy forms might be confused with P. longicollis but P. bolanderi can be distinguished by
its smaller size and dioicous inflorescences.
The leaves of P. bolanderi
are not only smaller but are relatively broader than those of P. longicollis. The var. seriata A. J. Shaw is characterized by whitish, ranked
leaves. It is not recognized here
because of intermediate plants from western 5.
Pohlia cruda (Hedwig)
H. Lindberg, Musci Scand., 18. 1879 Plants medium-size to robust, shiny green,
whitish-green, sometimes with a bluish tinge. Stems 0.5--2.5 cm. Leaves
0.8--2 mm, erect to ± spreading, sometimes slightly complanate, lanceolate to
elliptic, acute; margins serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells linear-rhomboidal, 70--140 \um,
vermicular, thin-walled. Specialized
asexual reproduction absent. Sexual
condition paroicous, rarely dioicous; perichaetial leaves strongly
differentiated, linear lanceolate, rarely weakly differentiated; perigonial
leaves in dioicous plants linear- or long-lanceolate from an ovate base. Seta orange to orange-brown. Capsule inclined 10--135° from
vertical, straw-colored to orange-brown, longly and slenderly pyriform, neck
ca. 1/2\x the urn length; exothecial
cells elongate-rectangular, walls straight; stomata superficial; operculum conic;
exostome teeth yellow-brown to red-brown, acute-triangular, pitted basally,
coarsely papillose distally; endostome hyaline, basal membrane 1/2\x the
exostome length, segments broadly keeled, broadly perforate, cilia short to
long. Spores 18--26 \um, ± coarsely
papillose. Capsules
mature summer (Jun--Aug). Soil banks, crevices in rocks or under roots,
tundra soil and paths; moderate to high elevations; Greenland; Alta., B.C.,
Man., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont., Que., Sask., Yukon;
Alaska, Ariz., Calif., Colo., Idaho, Iowa, Maine, Md., Mass., Mich., Minn.,
Mont., Nebr., Nev., N.J., N.Mex., N.Y., N.C., Oreg., Pa., S.Dak., Tenn.,
Utah, Vt., Va., Wash., W.Va., Wyo.; Mexico; South America; Eurasia; Pacific
Islands; Australia; Antarctic. Pohlia cruda is distinguished by glossy, pale green
to whitish or bluish leaves that are narrowly elliptic to lanceolate. The cells are long and narrowly
linear-vermicular. Perichaetial leaves are strongly differentiated, linear-lanceolate. Sporophytes have necks about as long as the
urn, with long-tapered exostome teeth, broadly keeled and perforate segments,
and short to long cilia. Along with P.
nutans, P. cruda is one of the most common North American species of the
genus. Unlike P. nutans, P. cruda is restricted to northern and montane sites
where it grows in rock crevices and on soil banks. The gametophytes can be unisexual
(dioicous) or bisexual (paroicous), a pattern of variation worthy of
additional study. 6.
Pohlia robertsonii J.
R. Shevock & A. J. Shaw, Bryologist 108: 177, fig. 1. 2005 Plants small to medium-size, dull green. Stems 0.5--1 cm. Leaves 1.3--1.6 mm, erect to ± spreading, lanceolate, acute;
margins serrulate to serrate in the distal 1/3; costa subpercurrent; distal
medial cells irregularly hexagonal to rhomboidal, 70--120 \um, thin- to firm-walled. Specialized asexual reproduction absent. Sexual condition paroicous, polysetous; perichaetial leaves
scarcely differentiated. Seta
orange-brown. Capsule inclined
10--45° from vertical, straw-colored to orange-brown, longly and slenderly
pyriform, neck ca. 3/4--1\x the urn length; exothecial cells
elongate-rectangular, walls slightly sinuose; stomata superficial; operculum
high-conic; exostome teeth orange-yellow to brown, bluntly acute, pitted
basally, coarsely papillose distally; endostome hyaline to yellow-brown,
basal membrane low, barely exceeding the capsule rim, segments narrow, keeled
at base, not or slightly perforate, cilia absent or rudimentary. Spores 16--20 \um, ± finely but
distinctly roughened. Capsules
mature spring (Mar-Apr). Sandy or gravelly soil, bases of sandstone or
volcanic rocks in open deciduous woodlands; low elevations; Pohlia robertsonii is the only North American polysetous
species of Pohlia. The leaves are less dull in appearance than
are those of P. elongata and P. nutans, but not notably shiny like
those of P. longicollis or P. cruda. They are more like the
moderately glossy leaves often seen in P.
bolanderi. The leaf cells have
firm but not thickened walls, thinner-walled than in P. elongata and P. nutans, shorter
and less vermicular than in P.
longicollis or P. cruda. Sporophytes are produced abundantly on the
paroicous gametophytes. The exostome
teeth are rather irregular and poorly developed, not gradually tapered to
narrow acute apices, and the endostome segments are narrow, scarcely keeled,
and narrowly or not at all perforate. 7. Pohlia obtusifolia (Villars
ex Bridel) L. Koch, Leafl. West.
Bot. 6: 20. 1950 Bryum obtusifolium Villars ex Bridel, Musc. Rec. 2(3):
52. 1803; P. cucullata
(Schwägrichen) Bruch ex Hochstetter Plants small to medium-size, dull pale green.
Stems 0.3--0.8 cm. Leaves 0.7--1.4 mm, erect to ± spreading,
broadly lanceolate, acute, subtly cucullate; margins serrulate to serrate in
the distal 1/3; costa subpercurrent; distal medial cells broadly rhombic,
30--70 \um, thin-walled and lax. Specialized asexual reproduction
absent. Sexual condition paroicous;
perichaetial leaves scarcely differentiated. Seta orange-brown. Capsule
inclined 160--180° from vertical, straw-colored to orange-brown, broadly
pyriform, neck about 1/3\x urn length; exothecial cells elongate-rectangular,
with straight walls; stomata superficial; operculum bluntly conic; exostome
teeth yellow to brown, rather slender, bluntly acute, weakly pitted basally,
coarsely papillose distally; endostome hyaline, basal membrane low, barely
exceeding the capsule rim to 1/3\x the exostome length, segments narrow,
delicate, weakly keeled, broadly perforate, cilia absent. Spores 17--25 \um, distinctly
roughened. Capsules mature summer (Jun--Aug).
Soil, often in late snowmelt areas in alpine and subalpine zones; high
elevations; Greenland; Alta., B.C., Nfld. and Labr., Nunavut, Ont., Que.,
Yukon; Alaska, Ariz., Calif., Colo., Maine, Mont., Nev., Tex., Vt., Wash.,
Wyo.; Eurasia. Plants of Pohlia obtusifolia with sporophytes
are generally small, but some sterile colonies in late snowmelt areas at high
elevations form deeper cushions. When
sterile, the subtly cucullate leaves provide a clue to their identity; they
are similar to those of P. drummondii
but the stems tend to be less red than in that species, the leaf cells are
broader and thinner walled, and axillary gemmae are absent. The plants are paroicous (unlike P. drummondii, which is dioicous) and
the capsules are barrel-shaped, with straight, rectangular walls (sinuose in P. drummondii and other gemmiferous
species). The peristome of P. obtusifolia is relatively reduced,
with slenderly triangular exostome teeth, a low endostomial basal membrane,
and narrow segments that are weakly keeled but broadly perforate. 8.
Pohlia crudoides (Sullivant
& Lesqereux) Brotherus in A. Engler & K. Prantl, Nat. Pfl. 1(3): 548.
1903 Bryum crudoides Sullivant & Lesquereux, Proc. Am.
Ac. Arts & Sc. 4: 278. 1859 Plants medium-size to robust, stiff, dull
green. Stems 0.4--2.5 cm. Leaves 1--2.2 mm, erect to ±
spreading, narrowly or rarely broadly lanceolate, acute; margins strongly
revolute, serrulate to serrate in the distal 1/3; costa subpercurrent; distal
medial cells linear-hexagonal, 35--80 \um, thin-walled. Specialized asexual reproduction absent. Sexual condition dioicous; perichaetial leaves moderately differentiated,
narrowly lanceolate; perigonial leaves shortly and broadly ovate. Seta orange-brown. Capsule inclined 30--90° from
vertical, brown, ovoid-cylindrical, neck about 1/3\x urn length; exothecial
cells shortly to longly rectangular, with straight or weakly sinuose walls;
stomata superficial; operculum conic; exostome teeth whitish to yellow,
weakly tapered, bluntly acute, coarsely papillose throughout; endostome
whitish to hyaline, basal membrane low, to 1/3\x the exostome length,
segments narrow, scarcely keeled, narrowly or not perforate, papillose, cilia
absent or rarely rudimentary. Spores
15--25 \um, finely roughened. Capsules mature summer (Jun--Aug).
Soil, generally in tundra, on banks and in depressions; low to high
elevations; Greenland; Alta., B.C., N.W.T., Nunavut, Que., Yukon; Alaska;
Eurasia. Pohlia crudoides is a distinctive Arctic species
characterized by cherry-red stems and relatively stiff, dark green leaves
with strongly recurved margins. The
perichaetial leaves of most Pohlia species
have recurved margins, but the only other North American species with
strongly recurved vegetative leaf margins is P. cardotii. The latter has shorter, broader leaves with
short-rhombic cells often with porose-thickened walls. Pohlia
crudoides has exostome teeth coarsely papillose from top to bottom, which
give them a whitish appearance under a dissecting microscope. 9. Pohlia ludwigii (Sprengel ex Schwägrichen) Brotherus,
Act. Soc. Sc. Fenn. 19: 27. 1892 Bryum ludwigii Schwägrichen, Spec. Musc. Frond.,
Suppl. 1(2): 95. 68. 1816 Plants medium-size to robust, dull green,
sometimes tinged with red. Stems
0.8--3.5 cm. Leaves 1.2--2.4 mm,
erect to ± spreading, lanceolate to broadly lanceolate, acute to bluntly
acute; serrulate to serrate in the distal 1/3; costa subpercurrent; distal
medial cells rhombic to rhomboidal, 40--80 \um, firm to rather thin-walled. Specialized asexual reproduction
absent. Sexual condition dioicous;
perichaetial leaves scarcely differentiated in shape but with costa very
strong at the base; perigonial leaves shortly and broadly ovate. Seta orange-brown. Capsule inclined 90--180° from
vertical, brown to straw-colored, broadly pyriform, neck about 1/3\x urn
length; exothecial cells shortly rectangular, with weakly to moderately
sinuose walls; stomata superficial; operculum bluntly conic; exostome teeth
yellow to brown, triangular-acute, pitted basally, coarsely papillose
distally; endostome hyaline, basal membrane to 1/2\x the exostome length, segments
distinctly keeled, broadly perforate, papillose, cilia short to long. Spores 14--21 \um, finely roughened. Capsules
mature summer (Jun--Aug). Soil, often in late snowmelt areas in alpine and
subalpine zones, in habitats similar to those of P. obtusifolia; high elevations; B.C., N.W.T., Ont., Que.;
Alaska, Ariz., Colo., Idaho, Mont., Wash.; Eurasia. Pohlia ludwigii is a rare alpine species characterized
by strongly decurrent leaves, often slightly cucullate apically. The older lower leaves generally have a
pink to reddish tinge. In late
snowmelt areas P. ludwigii can form
deep cushions when sterile. 10. Pohlia cardotii (Renauld in Renauld & Cardot)
Brotherus in A. Engler & K. Prantl., Nat. Pfl. 1(3): 547. 1903 Webera cardotii Renauld in Renauld & Cardot, Rev.
Bryol. 15: 71. 1888 Plants medium-size to robust, sometimes
forming deep turfs, dull green, sometimes tinged with red. Stems 0.4--2.5 cm. Leaves 0.7--1.5 mm, erect to ±
spreading, lanceolate to broadly lanceolate, acute; serrulate to serrate in
the distal 1/3; costa subpercurrent, very broad; distal medial cells rhombic,
18--50 \um, thick-walled, ± porose. Specialized
asexual reproduction absent. Sexual
condition dioicous; perichaetial leaves weakly differentiated,
elongate-lanceolate; perigonial leaves shortly and broadly ovate. Seta orange-brown. Capsule inclined 0--20° from
vertical, brown to straw-colored, narrowly pyriform, neck about 1/3\x urn
length; exothecial cells shortly rectangular, with sinuose walls; stomata
superficial; operculum bluntly to acutely conic; exostome teeth yellow to
brown, narrowly triangular-acute, pitted basally, coarsely papillose
distally; endostome hyaline, basal membrane scarcely exceeding the capsule
rim, segments narrow but weakly keeled, distinctly perforate, cilia absent to
rudimentary. Spores 14--19 \um,
finely roughened. Capsules mature summer (Jun--Aug).
Soil in mesic alpine and subalpine zones; high elevations; B.C.; A rare alpine
species, P. cardotii is
characterized by dull green, relatively broad leaves with a wide costa and
recurved margins. The sporophytes are
erect, with irregular exostome teeth, a low endostomial basal membrane, and
narrow, slightly keeled, broadly perforate segments. This species sometimes forms deep turfs in
moist alpine sites. 11. Pohlia
erecta Lindberg, Bot. Not. 1882: 195. 1882 Pohlia defecta
(Sanio) Andrews in Grout Plants slender, rather glossy green,
sometimes tinged with red. Stems
0.4--0.8 cm. Leaves 0.7--1.2 mm,
erect to ± spreading, lanceolate to broadly lanceolate, acute; serrulate to
serrate in the distal 1/3; costa subpercurrent; distal medial cells
rhomboidal, 35--60 \um, thin-walled. Specialized
asexual reproduction absent. Sexual
condition dioicous; perichaetial leaves weakly differentiated,
elongate-lanceolate; perigonial leaves shortly and broadly ovate. Seta orange-brown. Capsule inclined 0--20° from
vertical, brown to straw-colored, narrowly pyriform, neck ca. 1/3\x urn
length; exothecial cells shortly rectangular, with sinuose walls; stomata
superficial; operculum bluntly to acutely conic; exostome teeth yellow to
light brown, narrowly triangular, blunt, often irregular, pitted basally,
coarsely papillose distally; endostome hyaline, basal membrane scarcely
exceeding the capsule rim, segments rudimentary to elongate and weakly
keeled, ± perforate, cilia absent. Spores
15--21 \um, finely roughened. Capsules mature summer (Jun--Aug).
Soil in mesic alpine and subalpine zones; high elevations; B.C., N.W.T.; Pohlia erecta is a very rare species with soft,
sometimes reddish leaves having thin-walled cells. The erect sporophyte has a highly reduced
peristome with short, irregular exostome teeth and an endostome that consists
of a low basal membrane and poorly developed to rudimentary segments. |
|
12. Pohlia
lescuriana (Sullivant) Grout, Handlens. Microsc., 210. 1906 Bryum lescurianum Sullivant, Mem. Amer. Acad. Arts, n.s.
4: 171. 1849; Pohlia pulchella
(Hedwig) Lindberg Plants slender, pale whitish green. Stems 0.3--0.8 cm. Leaves 0.7--1.2 mm, erect to ±
spreading, lanceolate, acute, serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells linear-rhomboidal, 65--110 \um,
thin-walled. Specialized asexual reproduction
occasionally present; rhizoidal tubers orange, orange-brown, or yellow. Sexual condition dioicous;
perichaetial leaves weakly differentiated, lanceolate to linear-lanceolate;
perigonial leaves ovate, acuminate. Seta
orange-brown. Capsule inclined
95--180° from vertical, brown to straw-colored, shortly and rather broadly
pyriform, neck about 1/3\x urn length; exothecial cells short-rectangular,
with sinuose walls; stomata superficial; operculum low to high conic;
exostome teeth light yellow-brown, triangular-acute, pitted basally, coarsely
papillose distally; endostome hyaline, basal membrane 1/2\x the exostome
length, segments distinctly keeled, broadly perforate, cilia short to
rudimentary. Spores 15--21 \um,
finely roughened. Capsules
mature spring (Apr--Jun). On naturally or anthropogenically disturbed soils,
upturned tree banks, path banks, in crevices of rocks, along streams; low
elevations; N.B., Nfld. and Labr., N.S., P.E.I., Que.; Conn., Iowa, Maine,
Md., Mass., Mich., Mont., N.H., N.J., N.Y., N.C., Pa., Tenn., Vt., Va.,
W.Va.; Europe. Pohlia lescuriana is an inconspicuous woodland species
in eastern 13. Pohlia
pacifica A. J. Shaw, Contr. Plants slender, glossy green. Stems 0.3--1.0 cm, simple. Leaves 0.9--1.8 mm, erect to ±
spreading, lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells linear-vermicular, 110--145 \um,
thin-walled. Specialized asexual
reproduction absent. Sexual
condition dioicous; perichaetial leaves differentiated,
linear-lanceolate, to 3.2 mm; perigonial leaves ovate, elongate-acuminate. Seta orange-brown. Capsule inclined 120--180° from
vertical, brown to straw-colored, pyriform, neck about 1/3\x urn length;
exothecial cells short-rectangular, with sinuose walls; stomata superficial;
operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly keeled,
broadly perforate, cilia short to rudimentary. Spores 15--21 \um, finely roughened. Capsules
mature spring (Apr-Jun). Acid clay or sandy soils in disturbed places, stream
banks, ditches; low elevations; B.C., Pohlia 14.
Pohlia drummondii (Müller
Hal.) Andrews in Grout, Moss Fl. N. Am. 2: 196. 1935 Bryum drummondii Müller
Hal., Bot. Zeit. 20: 328. 1862; Pohlia
commutata Lindberg Plants medium-size, dark-green to
reddish-green. Stems 0.5--3.5 cm,
red. Leaves 0.9--1.5 mm, ±
spreading, lanceolate to ovate-lanceolate, acute; serrulate to serrate in the
distal 1/3; costa subpercurrent; distal medial cells rhombic to rhomboidal, 50--95
\um, thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising
singly in leaf axils, bulbiform, red (becoming ± black when dry), leaf
primordia at gemma apex and lower, laminate.
Sexual condition dioicous;
perichaetial leaves scarcely differentiated; perigonial leaves ovate, shortly
acuminate. Seta orange-brown. Capsule inclined 95--180° from
vertical, brown to straw-colored, pyriform, neck about 1/3\x urn length;
exothecial cells short-rectangular, with sinuose walls; stomata superficial;
operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules
mature summer (Jun--Aug). Acid, relatively humus-rich soils, alpine tundra,
stream banks, path banks; low to high elevations; Greenland; Alta., B.C.,
Nfld. and Labr., N.W.T., Nunavut, Ont., Que., Yukon; Alaska, Ariz., Calif.,
Colo., Idaho, Minn., Mont., Nev., N.H., N.Mex., Oreg., Utah, Vt., Wash.,
Wyo.; Europe. Pohlia drummondii has red stems (when moist) with cherry
red (when fresh) bulbiform gemmae in the leaf axils. The gemmae look like small branches, with
flexuose, laminate leaf primordia, and
occur single or occasionally in pairs in 1--5 distal leaf axils. The leaves pf P. drummondii are carinate and can be similar to those of P. obtusifolia, but are not cucullate
(and the plants are dioicous). 15.
Pohlia tundrae A.
J. Shaw, Bryologist 84: 65, fig. 1--10. 1981 Plants slender to medium-size, glossy, green
to light green. Stems 0.5--2.5 cm.
Leaves 0.9--1.5 mm, ± spreading,
lanceolate to ovate-lanceolate, acute; serrulate to serrate in the distal
1/3; costa subpercurrent; distal medial cells rhombic to rhomboidal, 45--95
\um, thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
clusters of 2--8 in leaf axils, narrowly branchlike to slender &
flexible, yellow, pink, or green, leaf primordia at apex and lower,
laminate. Sexual condition dioicous; perichaetial leaves scarcely
differentiated; perigonia leaves ovate, shortly acuminate. Seta orange-brown. Capsule inclined 95--180° from
vertical, brown to straw-colored, pyriform, neck about 1/3\x urn length;
exothecial cells short-rectangular, with sinuose walls; stomata superficial;
operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia short to rudimentary. Spores 16--23 \um, finely roughened. Capsules
mature summer (Jun--Aug). Acid, relatively humus-rich soils, alpine tundra,
stream banks, path banks, heavy metal mine tailings; low to high elevations;
Alaska, Calif., Colo., Idaho, Nev., N.Mex., Oreg., Utah, Wash., Wyo.; Europe. The gemmae of Pohlia tundrae are linear to oblong
and extend widely from the upper leaf axils.
They develop in clusters, sometimes densely. The leaf primordia are
broadly laminate and flexuose. Lax plants from shaded microsites have narrow
pale to whitish gemmae, but plants from more exposed sites have thicker,
sometimes reddish gemmae that can approach those of P. drummondii. At high
elevations in 16.
Pohlia rabunbaldensis A.
J. Shaw, Bryologist 108: 180, fig. 2. 2005 Plants slender, dull, green to light green. Stems 0.5--1 cm. Leaves 0.9--1.2 mm, ± spreading, lanceolate to ovate-lanceolate,
acute; serrulate to serrate in the distal 1/3; costa subpercurrent; distal
medial cells rhombic to rhomboidal, 45--95 \um, thin-walled. Specialized asexual reproduction
generally present when sterile; axillary gemmae arising singly in leaf axils,
narrowly bulbiform, orange to orange-red, leaf primordia at apex and lower,
sometimes to base, laminate, flexuose.
Sexual condition unknown,
presumed to be dioicous. Capsules
unknown. Acid, gravelly to sandy soil along trails; high elevations; Pohlia rabunbaldensis has bulbiform gemmae occurring singly
in a leaf axil, which are similar to those of P. drummondii, but the leaf primordia are even larger relative to
the gemma body. Plants of P. rabunbaldensis are slender, with
dull leaves much like those of P.
annotina, somewhat contorted when dry, distinct from the broader carinate
somewhat shiny leaves of P. drummondii,
which are unchanged on drying. 17. Pohlia filum
(Schimper) Mårtenssen, Svensk. Vet-Ak. Arh. Natur. 14: 149. 1956 Bryum filum Schimper,
1876, Syn. Musc. Eur. (ed. 2), 1876. Plants slender to medium-size, glossy, green
to light green. Stems 0.5--4 cm. Leaves 0.6--1.2 mm, erect, imbricate
to remote, lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to rhomboidal, 35--95 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae 1(--2) in
leaf axils, bulbiform, oblong or elliptic to subglobose, green to yellow,
becoming black when old, leaf primordia at apex, small, laminate. Sexual
condition dioicous; perichaetial leaves scarcely differentiated;
perigonia leaves ovate, shortly acuminate. Seta orange-brown. Capsule
inclined 95--180° from vertical, brown to straw-colored, pyriform, neck about
1/3\x urn length; exothecial cells short-rectangular, with sinuose walls;
stomata superficial; operculum convex-conic; exostome teeth yellow-brown,
narrowly triangular-acute, pitted basally, coarsely papillose distally;
endostome hyaline, basal membrane 1/2\x the exostome length, segments
distinctly keeled, broadly perforate, cilia short to rudimentary. Spores 16--23 \um, finely roughened. Capsules mature summer (Jun--Aug).
Gravely, organic-poor soils, glacial outwash, roadsides; low to high
elevations; Greenland; Alta., B.C., N.W.T., Nunavut, P.E.I., Que., Sask.;
Alaska, Oreg.; Europe. Pohlia filum is an easily recognized species
characterized by erect somewhat glossy leaves, and ovoid gemmae that normally
arise singly in the leaf axils. The
gemmae have a few small, triangular, rather stiff leaf primordia at the apex,
rarely with 1--2 primordia proximal on the gemma body. The gemmae of P. drummondii are more elongate-cylindrical and branch-like, with
larger, flexuose, often green leaf primordia at the apex and also frequently
more proximally on the gemma body. 18. Pohlia
beringiense A. J. Shaw, Contr. Plants slender, whitish green. Stems 0.5--2 cm. Leaves 0.9--1.5 mm, ± erect, lanceolate, acute; serrulate to
serrate in the distal 1/3; costa subpercurrent; distal medial cells
linear-rhomboidal, vermicular, 60--100 \um, thin-walled. Specialized asexual reproduction generally present; axillary
gemmae single in leaf axils, bulbiform, red (darker to black when dry), leaf
primordia at apex and more proximally, stiffly laminate. Sexual
condition unknown, presumed to be dioicous. Capsules
unknown. Acid, gravelly or sandy soil disturbed soil, ditches; low to high
elevations; Pohlia beringiense has cherry-red stems with erect,
whitish leaves and red bulbiform axillary gemmae. The gemmae are much like those of P. drummondii, but are even more red,
and contrast conspicuously with the whitish leaves. The organic-poor substrates that P. beringiense on which occurs are
quite different from the relatively humic soils on which P. drummondii occurs. 19.
Pohlia andalusica (Höhnel)
Brotherus in A. Engler & K. Prantl,
Nat. Pfl. 1(3). 551. 1903 Webera andalusica Höhnel, Sitzungsber. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Cl., Abt. 1, 104: 326. 1895. Plants slender to medium-size, ± glossy,
green. Stems 0.3--3 cm, red. Leaves 0.8--1.1 mm, ± erect,
lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to rhomboidal, 65--95 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
clusters of 2--8 in leaf axils, bulbiform, obconic to oblong, yellow, pink,
or green, leaf primordia mostly at apex, occasionally more proximally,
laminate. Sexual condition
dioicous; perichaetial leaves scarcely differentiated; perigonia leaves
ovate, shortly acuminate. Seta
orange-brown. Capsule inclined
95--180° from vertical, brown to straw-colored, pyriform, neck about 1/3\x
urn length; exothecial cells short-rectangular, with sinuose walls; stomata
superficial; operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules
mature summer (Jun--Aug). Acid, gravelly or sandy disturbed soil, path banks,
stream banks; Alta., B.C., N.B., Nfld. and Labr., N.W.T., N.S., Ont., P.E.I.,
Que., Yukon; Alaska, Calif., Colo., Idaho, Maine, Mass., Mont., N.Y., Pa.,
Vt., Wash., Wyo.; Europe. The gemmae of Pohlia andalusica occur in clusters of
2--6 in a few upper leaf axils and are reddish brown to green or sometimes
reddish-yellow. The gemmae are obconic
in shape, with laminate leaf primordia at the apices. This is one of the most common gemmiferous
species in New England, but is much more rare in western 20.
Pohlia bulbifera (Warnstorf)
Warnstorf, Krypt. Fl. Brandenburg 2: 429. 1904 Webera bulbifera
Warnstorf, Bot. Centrabl. 66:
230. 1896 Plants slender to medium-size, glossy, green.
Stems 0.3--3 cm, green, orange or
red. Leaves 0.9--1.4 mm, ± erect,
lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to rhomboidal, 65--95 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
clusters of 2--6 in leaf axils, spheroidal to obovate, yellow, orange, or
green, leaf primordia restricted to apex and forming a dome, laminate. Sexual condition dioicous;
perichaetial leaves scarcely differentiated; perigonia leaves ovate, shortly
acuminate. Seta orange-brown. Capsule inclined 95--180° from
vertical, brown to straw-colored, pyriform, neck about 1/3\x urn length;
exothecial cells short-rectangular, with sinuose walls; stomata superficial;
operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules
mature summer (Jun--Aug). Acid, gravelly or sandy disturbed soil, path banks,
stream banks, also on peaty soil; Greenland; B.C., Man., N.B., Nfld. and
Labr., N.W.T., N.S., Ont., Que.; Alaska, Colo., Maine, Mass., Mich., N.H.,
N.Y., Vt., Wash., Wis.; Eurasia. The gemmae of Pohlia bulbifera appear to be round to
oblong because the 4--5 laminate leaf primordia are concave and form a dome
over the gemma apex. The leaves are
wide-spreading so the gemmae, although small, can be seen clustered in the
distal leaf axils. This species sometimes occurs on peaty soils in bogs but
also occurs on gravelly soils in disturbed places. 21. Pohlia
annotina (Hedwig) Lindberg, Musci Scand., 17. 1879 Bryum annotinum Hedwig, Sp. Musc. Fond., 183. 1801 Plants slender, dull, green. Stems 0.3--3 cm, green, green to
orange-green. Leaves 0.6--1.1 mm,
± erect, lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to rhomboidal, 60--95 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
dense clusters in leaf axils, rarely few or single on older stems, oblong,
obconic, ± elongate and vermicular, rarely oblong-bulbiform, hyaline to pale
green, rarely reddish, leaf primordia 2--5, restricted to apex, peg-like or
rarely ± laminate with age. Sexual
condition dioicous; perichaetial leaves weakly differentiated, narrowly
lanceolate; perigonia leaves ovate, shortly acuminate. Seta orange-brown. Capsule
inclined 95--180° from vertical, brown to straw-colored, pyriform, neck about
1/3\x urn length; exothecial cells short-rectangular, with sinuose walls;
stomata superficial; operculum convex-conic; exostome teeth yellow-brown,
narrowly triangular-acute, pitted basally, coarsely papillose distally;
endostome hyaline, basal membrane 1/2\x the exostome length, segments
distinctly keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules mature summer (Jun--Aug).
On acid, gravelly or sandy disturbed soil, path banks, stream banks; B.C.,
N.B., Nfld. and Labr., N.S., Ont., Que.; Alaska, Ark., Calif., Del., Ga.,
Idaho, Iowa, Maine, Md., Mass., Mich., Miss., Mo., Mont., N.H., N.J., N.Y.,
N.C., Ohio, Pa., S.C., Tenn., Vt., Va., Wash., Wis.; Europe. Pohlia annotina is the most widespread and common
gemmiferous species in eastern 22.
Pohlia camptotrachela (Renauld
& Cardot) Brotherus, in A. Engler & K. Prantl, Nat. Pfl. 1(3): 552.
1903 Webera camptotrachela Renauld & Cardot, Bot. Gaz. 13(8):
199. 1888 Plants slender, dull, green. Stems 0.3--2.5 cm, green, green to
orange-green. Leaves 0.6--1.1 mm,
± erect, lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to rhomboidal, 60--95 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
dense clusters in leaf axils, globose to slightly elongate, yellow, green, or
orange, leaf primordia 1--4, restricted to apex, peg-like. Sexual condition dioicous;
perichaetial leaves weakly differentiated, narrowly lanceolate; perigonia
leaves ovate, shortly acuminate. Seta
orange-brown. Capsule inclined
95--180° from vertical, brown to straw-colored, pyriform, neck about 1/3\x
urn length; exothecial cells short-rectangular, with sinuose walls; stomata
superficial; operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules
mature summer (Jun--Aug). On acid, gravelly or sandy disturbed soil, path
banks, stream banks; B.C.; The leafy
stems of Pohlia camptotrachela are
indistinguishable from those of P.
annotina, but the gemmae are small, globose or subglobose, with 1--4
peg-like leaf primordial at the apex.
They are, in addition, translucent and may be yellow, green, or
orange. 23. Pohlia
proligera (Kindberg) Brotherus,
in A. Engler & K. Prantl, Nat. Pfl. 1(3): 551. 1903 Webera proligera Kindberg, Forh. Vidensk.-Selsk.
Kristiania 1888(6): 30. 1888 Plants medium-size, glossy, green. Stems 0.5--3.5 cm, green, green to
orange-green. Leaves 0.6--1.6 mm, ±
erect, lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to linear-rhomboidal, 60--105 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
dense felt-like clusters in leaf axils, oblong-linear to linear-vermicular,
hyaline, white, or pale green, rarely pale orange, leaf primordia 1--2(--3),
restricted to apex, peg-like. Sexual
condition dioicous; perichaetial leaves weakly differentiated, narrowly
lanceolate; perigonia leaves ovate, shortly acuminate. Seta orange-brown. Capsule
inclined 95--180° from vertical, brown to straw-colored, pyriform, neck about
1/3\x urn length; exothecial cells short-rectangular, with sinuose walls;
stomata superficial; operculum convex-conic; exostome teeth yellow-brown,
narrowly triangular-acute, pitted basally, coarsely papillose distally;
endostome hyaline, basal membrane 1/2\x the exostome length, segments
distinctly keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules
mature summer (Jun--Aug). Acid, sandy disturbed soil, path banks, stream
banks; low to high elevations. Greenland; Alta., B.C., N.B., Nfld. and Labr.,
N.W.T., N.S., Nunavut, Ont., P.E.I., Que.; Alaska, Ariz., Calif., Colo.,
Idaho, Maine, Mich., Minn., Mont., N.Y., Oreg., S.Dak., Utah, Vt., Wash.;
Europe. Pohlia proligera is a relatively common species in
boreal and high-montane habitats. The
leaves are glossy, with dense, felt-like clusters of whitish, pale green, or
sometimes pale orange gemmae in their axils.
The gemmae are linear-vermicular with 1--2 peg-like apical leaf
primordia. 24.
Pohlia andrewsii A.
J. Shaw, Bryologist 84: 70. 1981 Plants medium-size, glossy, green. Stems 0.5--2.5 cm, green, green to
orange-green. Leaves 0.6--1.6 mm,
± erect, lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to linear-rhomboidal, 60--105 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
dense clusters in leaf axils, isodiametric to oblong, green or more commonly
orange to red, leaf primordia 1--2(--4), restricted to apex, peg-like,
incurved over apex, becoming laminate with age. Sexual condition dioicous; perichaetial leaves weakly
differentiated, narrowly lanceolate; perigonia leaves ovate, shortly
acuminate. Seta orange-brown. Capsule inclined 95--180° from
vertical, brown to straw-colored, pyriform, neck about 1/3\x urn length;
exothecial cells short-rectangular, with sinuose walls; stomata superficial;
operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules
mature summer (Jun--Aug). On acid, sandy disturbed soil, path banks, stream
banks; low to high elevations. Greenland; N.W.T., The glossy
leaves of Pohlia andrewsii are much
like those of P. proligera but the
axillary gemmae are oblong, typically orange to red, and have 2--5 short,
primordia incurved over the gemma apex. 25. Pohlia flexuosa Harvey, Icon. Pl.
1: plate 19, fig. 5. 1836 Plants slender, dull, green. Stems 0.5--1.5 cm, green, green to
orange-green. Leaves 0.6--1.3 mm,
± erect, lanceolate, acute; serrulate to serrate in the distal 1/3; costa
subpercurrent; distal medial cells rhombic to linear-rhomboidal, 60--105 \um,
thin-walled. Specialized asexual
reproduction generally present when sterile; axillary gemmae arising in
dense clusters in leaf axils, dimorphic, isodiametric to oblong, brown,
knobby in outline, without leaf primordia, or narrowly linear-vermicular,
hyaline, with (0--)1--2 apical peg-like leaf primordia. Sexual condition dioicous; perichaetial leaves weakly
differentiated, narrowly lanceolate; perigonia leaves ovate, shortly
acuminate. Seta orange-brown. Capsule inclined 95--180° from vertical,
brown to straw-colored, pyriform, neck about 1/3\x urn length; exothecial
cells short-rectangular, with sinuose walls; stomata superficial; annulus
revoluble; operculum convex-conic; exostome teeth yellow-brown, narrowly
triangular-acute, pitted basally, coarsely papillose distally; endostome
hyaline, basal membrane 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia short to rudimentary. Spores 16--21 \um, finely roughened. Capsules
unknown in The slender,
dull plants of Pohlia flexuosa, which
have leaves somewhat contorted when dry, are like those of P. annotina and P. camptotrachela. The gemmae
are dimorphic, and are either hyaline to whitish and narrowly
linear-vermicular, with 1--2 poorly differentiated leaf primordia, or small,
blackish, oblong to globose, with angular cells. This species is known from only one North
American site but is common in parts of Asia including 26. Pohlia longibracteata Brotherus in J. Röll, Bot. Centralbl.
44: 419. 1890 Plants medium-size to robust, very glossy,
green to golden-green. Stems
0.5--2.5 cm. Leaves 2.0--3.5 mm,
wide-spreading, narrowly long-lanceolate, narrowly acute; serrate in the
distal 1/3; costa subpercurrent; distal medial cells linear-hexagonal,
80--160 \um, thin-walled. Specialized
asexual reproduction absent. Sexual
condition dioicous; perichaetial leaves ± strongly differentiated,
lanceolate to linear-lanceolate; perigonia generally on short lateral
branches; perigonial leaves ovate, very long-acuminate, to 6 mm. Seta orange-brown. Capsule inclined ± 180° from
vertical, brown to straw-colored, shortly pyriform to urceolate, neck less
than 1/3\x urn length; exothecial cells isodiametric, with straight to
sinuose walls; stomata immersed; annulus undifferentiated; operculum low to
high conic; exostome teeth light yellow-brown, triangular-acute, pitted
basally, coarsely papillose distally; endostome hyaline, basal membrane equal
to or slightly greater than 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia long, nodulose, or short. Spores 14--20 \um, finely roughened. Capsules
mature spring (May-Jul). On naturally or anthropogenically disturbed soils,
upturned tree banks, path banks, in crevices of rocks; along streams; low
elevations; B.C.; Alaska, Calif., Oreg., Wash. The relatively
long, narrow, wide-spreading leaves of Pohlia
longibracteata are quite glossy, easing identification of sterile
plants. The perigonial bracts are very
long-acuminate and spreading from the ovate base (providing the basis for the
specific epithet). Pohlia longibracteata occurs on soil
in Pacific coastal forests, typically under redwood trees. 27. Pohlia
brevinervis Lindberg & Arnell, Kongl. Svenska Vetenskapsakad. Handl. 23(10): 51. 1890. Plants slender to very slender, somewhat
glossy, whitish. Stems 0.3--1.5
cm. Leaves 0.6--1.4 mm, erect-spreading,
lanceolate, acute; serrulate in the distal 1/3; costa subpercurrent; distal
medial cells linear-rhomboidal, 60--110 \um, thin-walled. Specialized asexual reproduction
absent. Sexual condition dioicous;
perichaetial leaves weakly differentiated; perigonia leaves ovate, acuminate.
Seta orange-brown. Capsule inclined ± 180° from
vertical, brown to straw-colored, shortly pyriform to urceolate, neck less
than 1/3\x urn length; exothecial cells isodiametric, somewhat
collenchymatous, with sinuose walls; stomata superficial or slightly
immersed; annulus undifferentiated; operculum low to high conic; exostome
teeth dark to light brown or red-brown, triangular-acute, pitted basally,
coarsely papillose distally; endostome light yellow, basal membrane equal to
or slightly greater than 1/2\x the exostome length, segments distinctly
keeled, broadly perforate, cilia long, nodulose. Spores 16--21 \um, finely roughened. Capsules
mature spring (Apr-Jun). Clay soils along streams, ditches, glacial outwash;
Alta., Pohlia brevinervis is a rare and poorly known species that
seems to combine characters of P.
vexans (erect, glossy leaves on sterile plants) and P. atropurpurea (dark, reddish brown exostome teeth and somewhat
immersed stomates). The leaves are
distinctly more whitish than those of P.
vexans. 28. Pohlia columbica (Kindberg) Andrews in A. J. Grout, Moss
Fl. N. Am. 2: 202. 1935 Webera columbica Kindberg, Cat. Canad. Pl. 6: 115. 1892 Plants slender to medium-size, rather dull,
green to reddish-green. Stems
0.4--1.5 cm. Leaves 1.2--2.5 mm,
erect-spreading to spreading, narrowly lanceolate to lanceolate, acute;
serrate in the distal 1/3; costa subpercurrent; distal medial cells
linear-hexagonal, 85--150 \um, thin-walled. Specialized asexual reproduction absent. Sexual condition dioicous; perichaetial leaves ± differentiated,
to 3 mm, lanceolate to linear-lanceolate; perigonia leaves ovate,
elongate-acuminate, Seta
orange-brown. Capsule inclined
145--180° from vertical, brown to straw-colored, shortly pyriform to
urceolate, neck less than 1/3\x urn length; exothecial cells isodiametric,
with sinuose walls, somewhat collenchymatous; stomata immersed; annulus
undifferentiated; operculum low to high conic; exostome teeth yellow to
yellow-brown, triangular-acute, pitted basally, coarsely papillose distally;
endostome hyaline, basal membrane equal to or slightly greater than 1/2\x the
exostome length, segments distinctly keeled, broadly perforate, cilia long,
nodulose, or short. Spores 13--18
\um, finely roughened. Capsules
mature spring (May-Jul). Disturbed soils; along streams; low elevations;
B.C.; Pohlia columbica is a very rare and poorly known
species characterized by dull reddish plants, variably long-acuminate
perigonial bracts, pale yellow-brown exostome teeth, and immersed stomates. 29. Pohlia melanodon (Bridel) A. J. Shaw, Bryologist 84:
506. 1982 Bryum melanodon Bridel, Bryol. Univ. 1: 845. 1827; B. carneum Withering; B. delicatulum Hedwig Plants slender, soft and lax, dark green. Stems 0.3--1.0 cm. Leaves 0.7--1.3 mm, erect-spreading
to spreading, lanceolate to ovate-lanceolate, acute or shortly acuminate;
weakly serrulate in the distal 1/3; costa ending well before apex; distal medial
cells laxly and broadly rhomboidal, 60--100 \um, thin-walled. Specialized asexual reproduction
absent. Sexual condition dioicous;
perichaetial leaves scarcely differentiated, lanceolate; perigonia leaves
ovate, short-acuminate. Seta
orange-brown. Capsule inclined ±
180° from vertical, brown to red-brown, sometimes straw-colored, shortly
pyriform to urceolate, neck less than 1/3 urn length; exothecial cells
isodiametric, with sinuose walls, somewhat collenchymatous; stomata immersed;
annulus undifferentiated; operculum low to high conic; exostome teeth dark
brown to red-brown, triangular-acute, pitted basally, coarsely papillose
distally; endostome yellow to yellow-brown, basal membrane equal to or
slightly greater than 1/2\x the exostome length, segments distinctly keeled,
broadly perforate, cilia long, nodulose. Spores
13--18 \um, finely roughened. Capsules
mature spring (Apr-Jun). On naturally or anthropogenically disturbed clay or
rarely sandy soils; path banks, along streams; low elevations; Ont.; Pohlia melanodon has soft, dark green, rather broad
ovate-lanceolate leaves with lax, thin-walled cells. It does not produce elongate sterile
cushions as does P. wahlenbergii;
even when sterile the plants have short stems and are rather densely
leafy. The species is more common in
eastern 30. Pohlia atropurpurea (Wahlenberg) H. Lindberg, Acta Soc. F.
Fl. Fenn. 16: 14, figs. 1--7. 1899 Bryum pulchellum var. atropurpurea Wahlenberg, Fl. Lapp., 360. 1812 Plants very slender to tiny [[one reviewer says this are not
comparable descriptive terms]], soft, green to reddish. Stems 0.2--0.4 cm. Leaves 0.6--1.2 mm, erect-spreading
to spreading, lanceolate, acute; weakly serrulate in the distal 1/3; costa
ending well before apex; distal medial cells broadly rhomboidal, narrower in
perichaetial leaves, 65--110 \um, thin-walled. Specialized asexual reproduction absent. Sexual condition dioicous; perichaetial leaves scarcely
differentiated, lanceolate; perigonia leaves ovate, short-acuminate. Seta orange-brown. Capsule inclined ± 180° from
vertical, brown to red-brown, sometimes straw-colored, shortly pyriform to
urceolate, neck less than 1/3 urn length; exothecial cells isodiametric, with
sinuose, somewhat collenchymatous walls; stomata superficial; annulus
undifferentiated; operculum low to high conic; exostome teeth dark brown to
red-brown, triangular-acute, pitted basally, coarsely papillose distally;
endostome yellow to yellow-brown, basal membrane about 1/2\x the exostome
length, segments distinctly keeled, broadly perforate, cilia long, nodulose. Spores 15--21 \um, finely roughened. Capsules
mature spring (Apr-Jun). On naturally or anthropogenically disturbed clay or
rarely sandy soils; path banks, along streams; low elevations; B.C., Pohlia atropurpurea has sporophytes similar to those of P. melanodon, with dark reddish brown
exostome teeth, yellow endostomes, and immersed stomates. This uncommon northern species is very
small, generally consisting of unbranched gametophytes that do not form
extensive cushions. As in P. melanodon the leaf cells are lax
and thin-walled. Genetic relationships
between this and P. melanodon need
study; P. atropurpurea could be a
northern form of that species. Median
cells of the perichaetial leaves are narrower than the broadly lax cells of
proximal stem leaves. 31. Pohlia
vexans (Limpricht) H. Lindberg, Acta Soc. F. Fl. Fenn. 16:
20. 1899 Mniobryum
vexans Limpricht, Laubm. Deutsch. 2. 273. 1892 Plants slender, very glossy, green to
reddish. Stems 0.5--1.5 cm. Leaves 0.6--1.3 mm, stiffly erect and
± imbricate to erect-spreading, lanceolate, acute; weakly serrulate in the
distal 1/3; costa ending well before apex; distal medial cells broadly
rhomboidal, 65--110 \um, thin-walled. Specialized
asexual reproduction absent. Sexual
condition dioicous; perichaetial leaves somewhat differentiated,
lanceolate; perigonia leaves ovate, short- to ± long-acuminate. Seta orange-brown. Capsule inclined ± 180° from
vertical, brown to red-brown, sometimes straw-colored, shortly pyriform to
urceolate, neck less than 1/3 urn length; exothecial cells isodiametric, with
sinuose walls; stomata immersed; annulus undifferentiated; operculum low to
high conic; exostome teeth yellow to light brown, triangular-acute, pitted
basally, coarsely papillose distally; endostome yellow to yellow-brown, basal
membrane equal to or slightly greater than 1/2 the exostome length, segments
distinctly keeled, broadly perforate, cilia long, nodulose. Spores 15--21 \um, finely roughened. Capsules
mature spring (Apr-Jun). On naturally or anthropogenically disturbed clay or
rarely sandy soils; path banks, along streams; low to moderate elevations;
Alta., B.C., Man., N.W.T.; Alaska, Mont., Wash.; Europe. Pohlia vexans is a slender but handsome species
characterized by abundant, elongate sterile shoots with erect, rather glossy
leaves. The exostome teeth are pale
brown. The species is fairly common on
moist calcareous clays in cold continental regions of northwestern 32. Pohlia
wahlenbergii (F. Weber & D. Mohr) Andrews in A. J. Grout, Moss
Fl. N. Am. 2. 203. 1935 Hypnum wahlenbergii, Bot. Taschenb. 280. 475. 1807; Mniobryum wahlenbergii (F. Weber &
D. Mohr) Plants slender to very robust, dull, whitish
or sometimes in robust forms reddish. Stems
0.8--10 cm. Leaves 0.6--1.2 mm,
laxly spreading to more stiffly erect, lanceolate to ovate-lanceolate, acute;
weakly to strongly serrulate in the distal 1/3; costa ending well before
apex; distal medial cells broadly and laxly hexagonal to rhomboidal, 65--110
\um, thin-walled. Specialized asexual
reproduction absent. Sexual
condition dioicous; perichaetial leaves weakly differentiated,
lanceolate; perigonia leaves ovate, short-acuminate. Seta orange-brown. Capsule
inclined ± 180° from vertical, brown to red-brown, sometimes straw-colored,
shortly pyriform to urceolate, neck less than 1/3 urn length; exothecial
cells isodiametric, with sinuose walls, somewhat collenchymatous; stomata
immersed; annulus undifferentiated; operculum low to high conic; exostome
teeth light brown to red-brown, triangular-acute, pitted basally, coarsely
papillose distally; endostome hyaline to yellow, basal membrane equal to or
slightly greater than 1/2 the exostome length, segments distinctly keeled,
broadly perforate, cilia long, nodulose. Spores
15--21 \um, finely roughened. Capsules
mature spring (Apr--Jun). On naturally or anthropogenically disturbed clay or
rarely sandy soils; path banks, along streams; low elevations; Greenland;
Alta., B.C., Man., N.B., Nfld. and Labr., N.W.T., N.S., Nunavut, Ont.,
P.E.I., Que., Yukon; Alaska, Ariz., Ark., Calif., Colo., Conn., D.C., Idaho,
Ill, Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Mo., Mont.,
Nebr., Nev., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Oreg., Pa.,
S.Dak., Tenn., Utah, Vt., Va., Wash., W.Va., Wis., Wyo.; Mexico; Central
America; South America; Eurasia; Antarctica. Pohlia wahlenbergii is one of the most geographically
widespread and common species of the genus.
The leaves are pale whitish, decurrent, and have lax thin-walled
cells. The species varies greatly in
size, from very slender delicate plants in suboptimal habitats, to robust and
forming deep cushions in cold high altitude and latitude sites. Robust plants typically have red stems and
sometimes reddish pigmentation to the leaves as well. Some authors refer
robust expressions to the var. glaciale
(or P. glaciale) but gradation in
size and pigmentation is completely continuous, and different collectors draw
the line between the typical variety and the var. glaciale at different points. 3. Epipterygium Lindberg,
Öfvers. Förh. Kongl. Svenska
Vetensk.-Akad. 19: 603. 1862 * [Greek epi,
above, and pterigium, wing-like; alluding
to small dorsal leaves] Plants small to medium size, occurring in
loose turfs or as scattered plants. Stems
short to long, 0.1--1 cm, unbranched or irregularly forking when sterile,
often bearing subfloral innovations when with gametangia. Leaves spreading, weakly [strongly]
complanate, slightly [to very strongly] differentiated in 2--3 lateral and
ventral rows of larger leaves and 1--2[--3] dorsal rows of smaller narrower
leaves, 0.4--1.5[--2.5] mm, slightly decurrent; lateral and ventral leaves ovate-lanceolate
to elliptic, ± abruptly acute, dorsal
leaves smaller and narrower, serrulate near the apex or entire, costa ending
well before the apex; distal medial laminal cells broadly elongate-hexagonal
to long-rhomboidal, with thin to firm walls, marginal cells weakly but
distinctly [to strongly] differentiated from median cells. Specialized asexual reproduction
absent. Sexual condition dioicous;
perigonia and perichaetia terminal; perichaetial leaves somewhat
differentiated, lanceolate; perigonia bud-like, perigonial leaves ovate,
short- to ± long-acuminate. Seta
1, stramineous to pale orange, bent near the apex thus producing pendulous
capsules. Capsule inclined
160--180° from vertical, 1--3 mm, narrowly cylindric to short-pyriform or
urceolate, neck differentiated but short, stomata abundant in the neck,
superficial, median exothecial cells strongly collenchymatous, near mouth
quadrate or short-rectangular; annulus present and revoluble; peristome
diplolepideous-alternate; peristomial formula 4:2:6--8; exostome pale yellow
to brown or red-brown, teeth triangular, trabeculate, pitted basally,
coarsely or finely papillose distally, endostome hyaline to yellow, segments
wide, strongly keeled, broadly perforate, basal membrane high, cilia 1--3, as
long as the segments or shorter, nodulose or not. Species ca. 12
(1 in the flora); North America, Central America, South America, Europe,
Asia, Epipterygium is a small genus of tropical and temperate
mosses similar in morphology to Pohlia,
generally distinguished by more or less complanate foliation in which 2--3
rows of larger lateral leaves are differentiated in size and shape from 1--3
rows of smaller, narrower dorsal leaves.
The degree of leaf dimorphism reaches an extreme in E. wrightii of the Caribbean region,
but the only North American species, E.
tozeri, is almost uniformly foliate and scarcely complanate. This species is especially similar to Pohlia, and phylogenetic analyses
corroborate a close relationship between the two genera. SELECTED REFERENCES Shaw, A. J. 1984. Quantitative taxonomic study of morphology
in Epipterygium. Bryologist 87: 132--142. 1. Epipterygium
tozeri (Greville) Lindberg, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad.
21: 576. 1865. Bryum tozeri Greville, Scott. Crypt. Fl. 5: 285.
1827 Leaves loosely foliate and ± complanate,
weakly differentiated in 1--3 dorsal rows and 2--3 lateral andventral rows,
lateral and ventral leaves ovate-lanceolate to elliptic, ± abruptly acute, dorsal leaves smaller and
narrower, weakly serrulate in distal 1/3; costa ending well before apex;
distal medial cells broadly rhomboidal, 65--110 \um, 12--20 \um wide,
thin-walled. Seta orange-brown. Capsule with exostome teeth yellow to
light brown, triangular-acute, endostome yellow to yellow-brown, basal
membrane equal to or slightly greater than 1/2\x the exostome length,
segments distinctly keeled, broadly perforate, cilia long, nodulose. Spores 16--20 \um, finely roughened. Capsules mature spring (Apr-Jun).
On naturally or anthropogenically disturbed clay or more humus-rich soil,
wet, shaded banks along trails and streams; upturned tree bases; low elevations; B.C., Calif., Oreg., Wash.;
Europe; Asia. Epipterygium tozeri is a characteristic species of soil
banks in redwood forests of the Pacific coast. The soft pale bluish color is distinctive
when plants are growing in well-developed turfs. Plants of E. tozeri can be distinguished from any Pohlia by the (sometimes subtle) complanate foliation with
smaller dorsal than lateral and ventral leaves, broad, nearly elliptic
leaves, weak costa ending about 2/3 up the leaf, and by marginal leaf cells
differentiated from the median; longer, narrower, and more thick-walled. Illustrations,
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