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BFNA Title: Orthotrichum |
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Aug. 12, 2009
Second version ORTHOTRICHUM -
ORTHOTRICHACEAE 6. Orthotrichum
Hedwig, Spec. Musc. Frond., 162.
1801 * [For straight, erect hairs of calyptra in many species] Dale
H. Vitt Plants 1--5(--13) cm. Stem
leaves imbricate, usually erect-appressed and straight when dry, spreading
to wide-spreading when moist, 0.6--6.5 mm; ovate, lanceolate, or
linear-lanceolate, obtuse, acute to rounded-acute, acuminate or sometimes
obtuse, rarely channeled, rarely acuminate or piliferous; margins reflexed to
revolute, rarely plane or involute, entire or denticulate near apex; costa
strong, usually ending near apex; distal laminal cells mostly rounded, almost
isodiametric, usually 6--24(--30) \um, with 1--3 papillae per cell, papillae
conical or forked; basal leaf cells elongate-linear, thick-walled and nodose
or rectangular, thin-walled, and smooth near base; marginal leaf cells
shorter. Specialized asexual reproduction by brood-bodies,
sometimes present on leaves, 3--14 cells long. Perichaetial leaves similar to stem leaves or
sometimes slightly differentiated. Sexual condition goniautoicous or cladautoicous, rarely dioicous. Seta
to 15 mm, twisted to left. Capsule immersed, emergent or
exserted, varying from globose to cylindric, 0.7--3 mm, more or less
gradually constricted to seta, smooth or with 8 or 16 ribs and sometimes
constricted below mouth when dry; stomates superficial or immersed in central
or proximal parts of capsules; peristome double or more rarely single or
absent; exostome teeth 0, 8, or 16, papillose or striate, reflexed, recurved,
or erect; endostome segments 0, 8, or 16. Calyptra large, mitrate, conic-oblong to short-conic, more or
less hairy or naked, plicate or more rarely smooth, not split at base. Spores
isomorphic. Species 125
(47 in the flora); worldwide, more common in temperate areas. SELECTED
REFERENCES: Lewinsky, J. 1993.
Monographic studies on Orthotrichum
(Musci). Bryobrothera 2: 1--59. Vitt, D. H.
1973. A Revision of the Genus Orthotrichum in North America, North
of (For Key to
species, see end of treatment.) 1.
Orthotrichum affine
Bridel, Musc. Rec. 2(2): 22. 1801 Plants to 3 cm. Stem
leaves erect-appressed and straight
when dry, ligulate, elongate- to ovate-lanceolate, 2--4 mm, acuminate,
sharply acute, obtuse, or rounded, sometimes broadly apiculate; margins
recurved-revolute to just below apex, entire; distal laminal cells 9--13 \um
wide, 1-stratose with 2--3 low, forked papillae per cell; basal laminal cells
long-rectangular to elongate, nodose. Sexual condition goniautoicous. Seta
1--2.2 mm. Capsule 1/2 emergent to shortly exserted; urn oblong-cylindric
when mature, narrowly-cylindric and constricted along entire length when old
and dry, 1.6--2.5 mm, strongly 8-ribbed 2/3 entire length of capsule;
stomates superficial; peristome double; endostome segments 8, rarely 16, 2
cells wide at base, reticulate-papillose; exostome teeth 8, erect when
mature, reflexed-recurved when old, coarsely and obscurely papillose or
sometimes papillose-reticulate. Calyptra conic-oblong, with a few
papillose hairs. Spores 15--18 \um. Deciduous
trees, rarely on shaded rocks, shaded forests near stream beds and in
canyons, usually in mountainous areas; 0--3000 m; Alta., B.C.; Calif., Colo.,
Idaho, Mont., Oreg., Utah, Wash; Europe; Asia; n Africa. Orthotrichum affine, a species confined to western 2.
Orthotrichum alpestre
Hornschuch ex Bruch, Schimper & W. Gümbel, Bryol. Eur. 3 (fasc. 2/3) Orthotrichum
Suppl. 1: 1. 1849 Orthotrichum alpestre
var. majus Lesquereux & James; O. alpestre var. watsonii (James) Grout; O.
occidentale James; O. watsonii
James Plants 0.5--2.3 cm. Stem
leaves loosely erect-appressed and ± loosely twisted when dry; elongate-
to ovate-lanceolate, 2--3.5 mm, sharply acute, sometimes acuminate, cuspidate
or apiculate; margins revolute to below apex, entire; distal laminal cells
9--14 \um wide, 1-stratose, with 2--3 large, conical or forked papillae per
cell; basal laminal cells long-rectangular to rectangular, sometimes nodose
at insertion. Sexual condition goniautoicous.
Seta less than 1.4 mm. Capsule
1/2--3/4 emergent, sometimes almost exserted; urn oblong-ovate when mature,
oblong to short-cylindric when old and dry, less than 1.4 mm, strongly
8-ribbed 2/3 to the entire length; stomates immersed, just below middle of
the capsule, ± 1/2 covered by well-differentiated subsidiary cells; peristome
double; endostome segments 8 or 16, of 1 or 2 rows of cells, smooth or finely
roughened; exostome teeth 8, reflexed, densely and evenly papillose. Calyptra
oblong, ± hairy, hairs with large, distinct papillae. Spores
10--15 \um. Rocks and
trees, at the bases of trees and in moist crevices of large boulders,
especially common in moist, pine and deciduous forests; 2100--3000 m;
Greenland; Alta., B.C., Nfld., Ont., Yukon; Ariz., Calif., Colo., Idaho,
Mich., Mont., Nev., N.Mex., Oreg., S.Dak., Utah, Wash., Wyo.; Europe; Asia;
Pacific Islands (New Zealand). Orthotrichum alpestre
is distinguished by the highly papillose distal laminal cells, slightly
twisted, sharply pointed leaves, a calyptra with papillose hairs, the absence
of a preperistome, and the papillose or papillose-reticulate exostome. The endostome is always present in mature
capsules and, because of enlarged crosswalls, has a shiny appearance. The capsule is usually oblong and
strangulate, although the shape depends on age. In some specimens the capsule
is short-cylindric and not much constricted under the mouth. 3.
Orthotrichum anodon
Lara, Garilleti & Mazimpaka in Grarilleti, Lara, & Mazimpaka,
Bryologist 109: 188. 2006 Plants to 5 mm. Stem
leaves erect and slightly flexuose when dry, broadly ligulate to
ovate-ligulate, 2--2.5 mm, obtuse; margins recurved to near apex, entire;
distal laminal cells 10--16 \um wide, 1-stratose, smooth or with 1--2 low
papillae; basal laminal cells shortly rectangular, not nodose. Sexual
condition goniautoicous. Seta ± 0.5 mm. Capsule
immersed to shortly emergent, urn shortly cylindric to elliptic when dry,
1.4--1.6 mm, smooth and constricted at mouth by 8 short ribs; stomates
immersed in neck and proximal third of urn, completely covered by strongly
differentiated subsidiary cells; peristome single; endostome of 16 hyaline
and fragile, finely papillose segments; exostome none. Calyptra
conic, smooth, hairy. Spores 17--21 \um. Quercus trunks; low elevations; known from one
locality in Los Angeles Co., The small
plants, ligulate, blunt leaves, immersed stomates, lack of an exostome, and
capsule characters, unique among North American Orthotricha, distinguish Orthotrichum anodon. The puckered capsule mouth is reminiscent
of that of several species of Ulota
including U. obtusiuscula and U. coarctata. 4.
Orthotrichum anomalum
Hedwig, Spec. Musc. Frond., 162. 1801 Orthotrichum anomalum
var. americanum Macoun &
Kindberg; O. canadense Bruch &
Schimper; O. consimile ssp. anomaloides Kindberg Plants to .5 mm. Stem
leaves stiff, erect-appressed when dry, elongate-lanceolate, lanceolate,
or narrowly ovate-lanceolate, 2--3.8 mm, acute, bluntly acute, or narrowly
obtuse; margins recurved-revolute to near apex, smooth; distal laminal cells
9--14 \um wide, 1-stratose, with 1--3 small, conical papillae; basal laminal
cells long-rectangular to rectangular, sometimes slightly nodose. Sexual
condition goniautoicous. Seta 1--3 mm. Capsule
exserted; urn oblong-cylindric to cylindric, 1.8--2.4 mm, usually 16-ribbed
with 8 long ribs extending 1/2--2/3 length of capsule and alternating short 8
ribs; stomates immersed in middle and proximal portion of capsule; peristome
usually single; endostome rarely present; exostome teeth 16, rarely united
into 8 pairs, finely horizontally-striate below, vertically-striate
above. Calyptra oblong-conic, with few to many papillose hairs. Spores
11--15 \um. Rocks,
especially limestone, rarely at the base of trees or on dead branches;common
on the dryer rock faces in canyon areas; 0--3000 m; Greenland; Alta., B.C.,
Man., N.B., Nfld., N.W.T., N.S. Ont. Que., Sask., Yukon; Alaska, Ariz.,
Colo., Conn., Idaho, Ill., Ind., Iowa, Maine, Mich., Minn., Mont., N.Mex.,
N.Y., N.Dak., Ohio, Pa., R.I., S.C., S.Dak., Tenn., Utah, Vt., Va., Wash.,
Wisc., Wyo.; Mexico (Nuevo León, San Luis Potosí); West Indies (Haiti);
Europe; Asia; n Africa. Orthotrichum anomalum
has exserted capsules that are slightly constricted beneath the mouth, and
unistratose leaves. The most useful
character in distinguishing this species from others is the exserted capsule
with 8 long ribs alternating with 8 shorter ones. 5.
Orthotrichum bartramii
Williams, Bryologist 28: 76. 1925 Plants to 10 mm. Stem
leaves erect-appressed when dry, oblong to oblong-lanceolate, 2--3 mm,
bluntly cuspidate or sometimes irregularly obtuse or acute; margins recurved
to broadly revolute, incurved and serrate at apex; distal laminal cells 8--13
\um wide, 1-stratose, with two, small conical papillae; basal laminal cells
narrowly rectangular, grading to quadrate at margin, non-nodose. Sexual
condition goniautoicous or cladautoicous.
Seta less than 1 mm. Capsule
± emergent or rarely immersed; urn oblong to oblong-cylindric when old,
1.3--2 mm, 8-ribbed entire length; stomates immersed in proximal 1/2 of
capsule, 1/2 to completely covered by subsidiary cells; peristome double;
endostome segments 8, composed of 1--2 rows of cells, papillose, sometimes
coarsely so; exostome teeth 8, reflexed, densely and evenly papillose. Calyptra
oblong-conic, hairy, hairs papillose, deeply plicate. Spores
14--24 \um. Bark of Quercus or Populus tremuloides; ±
3000 m; Ariz.; Mexico; Central America (Guatemala). The channeled,
cuspidate apex of the mature leaves distinguishes Orthotrichum bartramii from all other species with immersed
stomates. The sporophytic characters
strongly resemble those of O. tenellum;
however, the latter species has non-channeled, incurved leaf tips 6.
Orthotrichum bolanderi
Sullivant, Icon. Musc. Suppl., 64, plate 46.
1874 Plants 0.5--4 cm, dark-green to black. Stem
leaves stiffly erect-appressed,
rarely incurved when dry, lanceolate, 2--3.5 mm, narrowly obtuse to acute;
margins plane, entire; distal laminal cells 7--11 \um wide, 2-stratose, each
cell with 2--4 conical, sometimes large papillae; basal laminal cells
elongate, with ± wavy walls, smooth. Sexual condition goniautoicous. Seta
± 1 mm. Capsule emergent; urn ovate-oblong when mature, ovate-cylindric
when old, 1.4--2 mm, lightly 8-ribbed in distal 1/3 of capsule; stomates
superficial; peristome double; endostome segments 8, 2 rows of cells,
papillose; exostome teeth 8, reflexed-recurved, papillose to coarsely
papillose-striate. Calyptra oblong-conic, with papillose
hairs. Spores 17--23 \um. Rocks,
especially sandstone; 100--2000 m; Orthotrichum bolanderi
is the only species with superficial stomates that has 2-stratose distal
laminal cells. It has leaves with subsheathing bases, ovoid cylindric
capsules with long wrinkled necks, and reflexed-recurved exostome teeth. When dry, the leaves are stiffly
erect-appressed and they lie flat on a microscope slide when wet. 7.
Orthotrichum consimile
Mitten, J. Linn. Soc. Bot. 8: 24. 1864 Orthotrichum columbicum
Mitten; O. hendersonii Renault & Cardot; O. pulchellum var. columbicum
(Mitten) Grout; O. pulchellum var. leucodon Venturi; O. pulchellum var. longipes
Lesquereux & James; O. pulchellum
var. productipes Renault &
Cardot; O. pulchellum ssp. ulotaeforme (Renault & Cardot)
Kindberg; O. ulotaeforme Renault
& Cardot Plants 8--20 mm. Stem
leaves erect and loosely flexuose to crisped-flexuose when dry,
linear-lanceolate, 2--3.2 mm, acuminate (sometimes bluntly so) or acute;
margins recurved to near apex, sometimes reflexed above, entire; distal
laminal cells 7--11 \um wide, 1-stratose, smooth or bluntly papillose; basal
laminal cells linear-elongate to long-rectangular, non-nodose. Sexual
condition goniautoicous. Seta (1.5--)2.5--6 mm. Capsule
exserted to long-exserted; urn ovate-oblong when young, elliptic-cylindric
when old, 2.5--6 mm, 8-ribbed entire length; stomates numerous, immersed, in
neck, proximal, and middle portion of capsule 1/3--2/3 covered by strongly
differentiated subsidiary cells; peristome double; endostome segments usually
16, rarely 8 (sometimes 8 long and 8 short), of 1 row of cells, smooth or
finely roughened; exostome teeth 8, erect when mature, reflexed when old,
very finely and densely papillose on outside, finely striate on inner
surface. Calyptra oblong-conic to short-conic, naked or rarely sparsely
hairy. Spores 12--14 \um. Tree trunks
and branches in coniferous forests, sometimes found on boulders in shaded
habitats; low elevations; B.C.; Orthotrichum consimile
is distinguished from superficially similar species of Ulota by its glabrous calyptra, immersed stomates, and
undifferentiated basal leaf cells.
From O. pulchellum, it differs by having stomates confined to the
proximal portion of the capsule and an exostome of 8 yellowish teeth that are
finely papillose on the outer surface and finely striate on the inner. Also, O.
consimile is much larger and has
capsules that are long-exserted and constricted beneath the mouth when old
and dry. 8.
Orthotrichum cupulatum
Bridel, Musc. Rec. 2(2): 25. (May 9) 1801 Orthotrichum leiodon
Kindberg; O. utahense Lesquereux Plants to 12 mm. Stem
leaves erect-appressed when dry, ovate-lanceolate to oblong-lanceolate,
2.5--4 mm, acute or rarely narrowly obtuse; margins recurved or revolute to
near the apex, entire; distal laminal cells 8--12 \um wide, 1-stratose, or
occasionally areas with 2-stratose, densely pluri-papillose with 2--3 large,
forked papillae or papillae sometimes small and conical; basal laminal cells
long- to short-rectangular, non-nodose.
Sexual condition
goniautoicous. Seta 0.5--1 mm. Capsule immersed to 1/2 emergent when
dry; urn ovate-pyriform to sometimes short-oblong, 1.2--1.6 mm, ± 0.8 mm
wide, 16-ribbed; stomates immersed, in 1 or 2 rows in the middle of the
capsule; peristome single, rarely double; endostome segments rarely present,
8, 1 row of cells wide, smooth; exostome teeth 16, erect, becoming
spreading-reflexed with age, usually not present when the capsule is old,
papillose-striate or papillose-reticulate.
Calyptra oblong to
oblong-conic, with papillose hairs. Spores 9--16 \um. Faces of
calcareous cliffs and large boulders in coniferous forests, most common in
the dryer areas of Ponderosa Pine forests and on the dryer cliff faces in
canyons; 100--3000 m; B.C.; Ariz., Calif., Colo., Idaho, Mont., Nev., Oreg.,
S.Dak. Texas, Utah, Wash., Wyo.; Mexico (Nuevo León). Europe; Asia; n Orthotrichum cupulatum
is extremely variable but can be distinguished by the following combination
of characters most of which are usually evident: (1)
leaves ovate-lanceolate, broadly but sharply acute, and usually
unistratose or occasionally partially 2-stratose; (2)
papillae of the leaf cells usually large and forked, but if small and
conical then the leaf is unistratose;
(3) margin of leaf never
thicker than lamina; (4) peristome
teeth always 16, erect or spreading, and yellow-brown; and (5) capsule
16-ribbed, sometimes with 8 short and 8 long ribs. 9.
Orthotrichum diaphanum
Bridel, Musc. Rec. 2(2): 29. 1801 Orthotrichum garretii
Grout & Flowers Plants 2--7 mm. Stem
leaves loosely-erect when dry, ovate-lanceolate to oblong, 1.8--3 mm,
gradually acute to a ± serrate, hyaline awn; margins revolute nearly to base
of awn; distal laminal cells 14--24 \um wide, 1-stratose, smooth or with 1-2
small, conical papillae; basal laminal cells broadly rectangular, non
nodose. Sexual condition autoicous.
Seta less than 1 mm. Capsule
immersed to emergent; urn oblong to oblong-cylindric, 1.2--1.8 mm,
wrinkled or moderately 8-ribbed when dry; stomates immersed, numerous, at
middle of capsule; peristome double; endostome segments 16, of 1 or 2 rows of
cells, densely papillose; exostome teeth 16, reflexed when old and dry,
longitudinally-striate or coarsely and densely papillose. Calyptra
conic-oblong, naked or with a few, smooth hairs. Spores
13--20 \um. Bases and
trunks of trees less than 2 m, in dry, usually exposed areas, rarely on rock;
100--1000 m; B.C.; Alaska, Ariz., Ark., Calif., Colo., Kans., La., Mo., Neb.,
Ohio, Okla, N.Mex. S.Dak., Tenn., Texas, Utah; South America (Uruguay) ;
Europe; n Asia; Africa (Algeria); Atlantic Islands (Canary Islands). Orthotrichum diaphanum
is distinguished by the hyaline awn of the distal stem leaves. Gametophytically, O. diaphanum is most
closely related to O. pumilum, with which it shares the
following attributes: large
chlorophyllose leaf cells lacking large papillae, presence of numerous
gemmae, and ovate-lanceolate leaves. Orthotrichum
pumilum has apiculate leaf apices
that often are similar to those found on the young plants of O. diaphanum. 10.
Orthotrichum elegans Hooker & Greville, Edin. Jour. of Sc. 1 (1): 122. 1824 Orthotrichum psilothecium
Macoun & Kindberg; O. speciosum
ssp. elegans (Hooker &
Greville) Lindberg; O. speciosum
var. elegans (Hooker &
Greville) Warnstorf; O. speciosum
var. roellii Venturi Plants 5--22 mm. Stem
leaves stiffly flexuose when dry, wide-spreading when moist, lanceolate
to narrowly lanceolate, 2.3--3 mm, broadly acuminate to acute, margins
strongly revolute nearly to apex, entire; distal laminal cells 8--10 \um
wide, 1-stratose, with small conical papillae; basal laminal cells elongate,
nodose. Sexual condition autoicous.
Seta 1--2 mm. Capsule
smooth, erect, emergent to sometimes shortly exserted; urn small and narrow,
narrowly cylindric to fusiform-cylindric, 1.3--2 mm, smooth; stomates
superficial; peristome double; endostome segments 8, 2 rows of cells,
coarsely papillose; exostome teeth 8, recurved, coarsely papillose. Calyptra
conic-oblong, sparsely hairy. Spores 13--26 \um. Deciduous trees, especially Populus species; occasionally conifers; 200--1000 m; Alta., B.C., Man., N.B., Nfld., N.W.T., N.S., Ont., P.E.I., Que., Sask.; Alaska, Maine, Mass., Mich., Minn., Mo., N.H., N.J., N.Y., Ohio, Pa., R.I., Vt., Wis.; Europe; Asia. The small plants of Orthotrichum elegans in a rather compact tuft are quite different from the larger, loosely tufted plants of O. speciosum. Orthotrichum elegans also has fusiform-cylindric, totally smooth, and emergent to shortly exserted capsules while O. speciosum has long-cylindric, generally lightly 8-ribbed, fully exserted capsules. Orthotrichum striatum has 16 exostome teeth and endostome segments. 11.
Orthotrichum euryphyllum
Venturi, Bot. Centralbl. 44: 418. 1890. Orthotrichum epapillosum E. Lawton & F. J. Hermann Plants to 3 cm, in dense, dark red-brown to
black tufts. Stem leaves appressed and erect with reflexed apices when dry,
oblong-lanceolate to ovate-lanceolate, 2.5--4 mm, obtuse and cucullate at
apex, margins recurved-revolute to near apex, entire or sometimes irregularly
serrulate near apex; distal laminal cells to 20 \um wide, 1-stratose, smooth
or with 1--2 low papillae; basal laminal cells short-rectangular with
somewhat nodose walls. Sexual condition goniautoicous. Seta
to 1.5 mm. Capsule immersed, urn oblong to oblong-ovate, 1.5--2 mm long,
strongly 8-ribbed the entire length; stomates immersed in proximal part of
capsule, completely covered by subsidiary cells; peristome double; endostome
segments (8--)16, well-developed, persistent, finely papillose; exostome
teeth 8 splitting to 16, erect until old, then loosely reflexed, densely
papillose-striate. Calyptra 16-plicate with long beak,
naked. Spores to 15 \um. Calcareous
springs in Artemisia scrub; 0--500
m; Orthotrichum euryphyllum has compact, red-brown plants,
cucullate, entire leaf apices, and distal laminal cells approaching 20 \um in
width. In comparison, O. rivulare has loose, blackish
plants; flat, dentate leaf apices, and small distal laminal cells rarely
wider than 13 \um. 12.
Orthotrichum exiguum
Sullivant in A. Gray, Man. Bot. No. U. States ed. 2: 633. 1858 Plants 3--10 mm. Stem
leaves erect-appressed when dry, lanceolate-oblong to oblong, 0.7--1.3
mm, rounded-obtuse to abruptly tapering to a broadly acute apex; margins
reflexed to broadly recurved, entire; distal laminal cells 7--13 \um wide,
1-stratose, with conical or c-shaped papillae; basal laminal cells
irregularly rounded. Sexual condition goniautoicous. Seta
less than 5 mm. Capsule immersed or slightly emergent, urn ovate-oblong, less
than 1 mm, distinctly 8-ribbed to the base when dry; peristome double; endostome
segments 8, as wide as exostome teeth, granulate-papillose; exostome teeth
16, spreading or reflexed when dry, papillose to cross-striolate below,
papillose to vertically or reticulately striolate above. Calyptra
short-conic, naked or with a few hairs near tip. Spores
17--20 \um. Base of trees
or on tree trunks; 10--200 m; The distal
laminal cells with bulging walls and c-shaped papillae, delicate, hyaline
endostome, clavate stems, papillose, quadrate basal cells, differentiated
perichaetial leaves, and capsules that are widest at the mouth, with 16,
reflexed exostome teeth, separate this species from all others. 13.
Orthotrichum fenestratum
Cardot & Thériot, Proc. Wash. Ac. Sc. 4: 310. 16. 1902 Plants to 12 mm. Stem
leaves erect-appressed when dry,
ovate-lanceolate to narrowly lanceolate, 2.2--3.2 mm, acute and often
cuspidate-apiculate; margins narrowly revolute to below apex, entire; distal
laminal cells 9--15 \um wide, 1-stratose, with 1--2 small conical papillae
per cell; basal laminal cells narrowly rectangular, grading to subquadrate at
margin, nodose, thick-walled. Sexual condition goniautoicous. Seta
± 2 mm. Capsule exserted; urn ovate when moist, ovate-oblong when dry,
1--1.8 mm, smooth to moderately 8-ribbed; stomates superficial; peristome
single, rarely double; endostome segments 8, narrow, consisting of 1 row of
cells, papillose-reticulate; exostome teeth 8 splitting to 16, erect to
recurved when old, coarsely papillose distally, becoming finely papillose to
almost smooth near base, fenestrate and cancellate. Calyptra
short-conic, plicate, with few, thick, white hairs. Spores
20--27 \um. Collected only
once, by J. M. Macoun, who did not record the substrate; probably near sea
level; The exostome
teeth that are cancellate to the base, and fenestrate and perforate only in
the distal portion, and the ovate capsule, distinguish Orthotrichum fenestratum. 14.
Orthotrichum flowersii
Vitt, Bryologist 74: 159. 1971 Plants to 5 mm. Stem
leaves stiffly-erect and loosely-appressed when dry, ovate-oblong to
oblong-ligulate, 2--2.8 mm, rounded or obtuse, often mucronate, carinate,
subcucullate; margins revolute, entire or rough by means of projecting papillae near apex;
distal laminal cells 13--19 \um wide, 1-stratose, with ± 2, small, conical
papillae; basal laminal cells rectangular to short-rectangular,
non-nodose. Sexual condition autoicous.
Seta ± 0.5 mm. Capsule
immersed or shortly emergent; urn ovate-oblong to oblong when mature,
1--1.4 mm, strongly 8-ribbed 1/2 to 3/4 the length of the capsule, stomates
immersed, in 1 or 2 rows at middle of the capsule, guard cells barely to 1/2
covered by subsidiary cells which have inner walls ± thickened; peristome
double; endostome segments 8, usually of 1 row of cells, finely roughened;
exostome teeth 8, reflexed, densely papillose or papillose-striate
above. Calyptra oblong, naked or with a few hairs. Spores
10--17 \um. Base of tree
trunks and trunks of deciduous trees in xeric and exposed habitats; 500--2000 m; Orthotrichum flowersii is distinguished from O. pumilum
and O. pallens by blunt, carinate leaves that are ovate or ovate-oblong,
almost always with a small mucro. 15. Orthotrichum gymnostomum Bridel, Bryol. Univ. 1: 782. 1827 Nyholmiella gymnostoma
(Bridel) Holmen & E. Warncke Plants 4--12 mm. Stem
leaves imbricate, erect-appressed when dry, ovate to ovate-oblong, concave,
2--2.8 mm; rounded to obtuse; margins involute, entire; distal laminal cells
7--12 \um wide, 1-stratose, with 2--3 conical papillae; basal laminal cells
elongate-rectangular, non-nodose. Sexual condition dioicous. Seta
± 0.5 mm. Capsule immersed; urn oblong, 1.8 mm; moderately 8-ribbed when
dry; stomates superficial, in one row at distal end of capsule; peristome
none. Calyptra obscurely plicate, naked, papillose. Spores
18--20 \um. Bark of
deciduous trees; 0--200 m; Nfld.; Europe; Asia ( Orthotrichum gymnostomum differs from the similar O. obtusifolium
by leaves with involute margins, laminal cells with 2--3 papillae, and
immersed capsules with no peristome, while O. obtusifolium has
leaves with erect margins, 1-papillose cells, and emergent capsules with a double peristome. Orthotrichum gymnostomum is rare in North America, restricted to the coastal
areas of 16.
Orthotrichum hallii
Sullivant & Lesquereux in W. S. Sullivant, Icones Musc. Suppl., 63, plate 45.
1874 Plants to 2.5 cm. Stem
leaves loosely erect-appressed when dry, elongate-lanceolate to
lanceolate, 1.7--3.5 mm, ± narrowly obtuse, rarely bluntly acute; margins
revolute below, recurved to almost plane near apex, entire; distal laminal
cells 7--13 \um wide, 2-stratose, rarely with a few unistratose areas,
papillae always small, conical, 1--3 per cell; basal laminal cells
rectangular to short-rectangular, grading to quadrate at margins,
non-nodose. Sexual condition goniautoicous.
Seta 0.5--1 mm. Capsule
immersed when moist, about 1/2 emergent when dry, urn oblong, short-oblong,
or oblong-ovate, 1--1.8 mm, strongly 8-ribbed 1/2 to entire length, rarely
with 8 very short intermediate ribs; stomates immersed; peristome double;
endostome segments 8, of 1 row of cells, finely longitudinally-striate;
exostome teeth 8, sometimes irregularly split to 16, incurved until old, then
spreading or rarely reflexed, coarsely striate-reticulate, or
striate-papillose. Calyptra oblong, sparsely hairy,
hairs papillose. Spores 10--17 \um. Rocks, usually
limestone or calcareous sandstone, sometimes granite, quartzite, or basalt,
rarely on trunks of deciduous trees, open pine forest, spruce-fir forests or
deciduous scrub oak-maple forests, especially common on vertical canyon walls
and shaded cliff faces; 200--3,000 m; B.C.; Ariz., Calif., Colo., Idaho,
Mont., Nev., N.Mex., S.Dak., Utah, Wyo.
Restricted to western Orthotrichum hallii
is distinguished by having leaves that are completely, or nearly 2-stratose
in their distal portion, and emergent, oblong, 8-ribbed capsules. The peristome teeth vary from
ridged-striate to papillose-striate. The proximal portions of the teeth are
often coarsely papillose with the distal portions having striae mixed with
papillae. The papillae are always small and conical. 17.
Orthotrichum holzingeri
Renault & Cardot in J. M. Holzinger, Contr. U.S. Nat. Herb. 3: 270. 1895 Plants to 3 cm. Stem
leaves erect, loosely-appressed
when dry, oblong-lanceolate to ovate-lanceolate, 1.8--3 mm, broadly acute to
obtuse; margins narrowly recurved-revolute to near apex, entire; distal
laminal cells 7--12 \um wide, 1-stratose, smooth; basal laminal cells
rectangular, not nodose in small leaves, long-rectangular and sometimes
nodose in larger leaves. Sexual condition goniautoicous, perigonia numerous. Seta
2--3 mm. Capsule fully exserted; urn cylindric, 1.5--2.2 mm, slightly to
distinctly 8-plicate for 1/2 length; stomates superficial; peristome single;
endostome segments absent; exostome teeth 16, erect to spreading-reflexed
when old and dry, smooth. Calyptra short-conic, naked. Spores
13--18 \um. Seasonally wet
rocks in small streams of dry montane forests; 1000--2000 m; Orthotrichum holzingeri is distinguished by smooth leaf cells,
smooth exostome, naked calyptra, and exserted, lightly-ribbed capsules. It is a distinct species, endemic to a
small region in western 18.
Orthotrichum keeverae
H. A. Crum & L. E. Anderson, J. Elisha Mitchell Sc. Soc. 72: 281. 1956
Plants 0.5--2 mm. Stem
leaves rigid, erect-appressed when
dry, oblong-lanceolate from ovate base to narrowly lanceolate, 2.2--4 mm, slenderly
acute, tip contracted to a large, cuspidate point; margins revolute nearly to
apex, often irregularly notched near apex; distal laminal cells 7--14 \um
wide, 1-stratose, with small, conical papillae, basal laminal cells
linear-elongate, nodose. Sexual condition goniautoicous. Seta
± 1 mm. Capsule about 1/2 emergent when dry; urn cylindric, 1.5--2 mm,
distinctly 8-ribbed entire length when mature; stomates superficial;
peristome double, endostome segments 16, almost as wide as exostome teeth, densely
papillose; exostome teeth 16, recurved when dry, densely papillose, often
united at base. Calyptra conic-oblong, with papillose hairs 1--3 cells
thick. Spores 17--27 \um. Trunks and
branches of live and dead Quercus Orthotrichum keeverae
is the only species in southeastern 19.
Orthotrichum kellmanii
Norris, Shevock & Goffinet, Bryologist 107: 210. 2004 Plants to 10 mm. Stem
leaves erect-appressed when dry, ovate-ligulate to ovate-oblong, grading
to lanceolate or ovate lanceolate on reproductive stems; 1.4--2.5 mm, obtuse,
acute to broadly acuminate near perichaetia; margins recurved to near apex,
entire; distal laminal cells 10--12 \um wide, uniformly 2-stratose, with low,
simple papillae; basal laminal cells quadrate to short rectangular;
non-nodose. Sexual condition goniautoicous.
Seta ±1 mm. Capsule
immersed; urn ovate-oblong, ± 1.5 mm, 8-ribbed entire length; stomates
immersed in proximal half of capsule, mostly covered by subsidiary cells;
peristome double; endostome segments 8, delicate and smooth, fragile; exostome
teeth 8, blunt, cribose and splitting to 16, densely and coarsely papillose,
erect to spreading when dry. Calyptra short-conic, with sparse,
papillose hairs. Spores 8--12 \um. Dry sandstone
boulders in dense chaparral; Only two
species of Orthotrichum have
immersed stomates and uniformly bistratose distal laminal cells. Orthotrichum
hallii is a species of calcareous rocks found sporadically throughout the
west, while O. kellmanii occurs on
sandstone boulders only at a few sites in central 20.
Orthotrichum laevigatum
Zetterstedt, Oefv. Svensk. Vet. Ak. Foerh. 19: 363. (May 14) 1862 Orthotrichum hainesiae
Austin; O. kingianum Lesquereux; O. laevigatum var. kingianum (Lesquereux) Grout; O. lonchothecium Macoun &
Kindberg; O. macounii Austin; O. macounii var. lonchothecium (Macoun & Kindberg) Grout; O. raui Austin; O. roellii
Venturi; O. roellii ssp. schlotthaueri (Venturi) Kindberg; O. roellii ssp. stenocarpum (Venturi) Kindberg; O. schlotthaueri Venturi; O.
speciosum var. brevicaule
Lesquereux; O. speciosum var. hainesiae (Austin) Paris; O. speciosum ssp. laevigatum (J. E. Zetterstedt) J. J. Amann; O. speciosum var. polyanthum
Lesquereux; O. speciosum var. polycarpum Lesquereux & James; O. speciosum var. raui (Austin) Lesquereux & James Plants 0.5--4.3 cm, usually ± 2 cm. Stem
leaves erect-appressed, slightly secund and stiff when dry,
oblong-lanceolate to lanceolate to ovate-lanceolate, 2--4.3 mm, obtuse,
bluntly acute, or gradually acuminate to narrowly acute; margins revolute
from base to near apex, entire; distal laminal cells 8--12 \um wide,
1-stratose, densely papillose with low, 1--3 conical or low-forked papillae;
basal laminal cells rectangular to elongate, grading to elliptic-quadrate,
nodose. Sexual condition
goniautoicous. Seta 1.5--4 mm. Capsule fully exserted; urn
cylindric, 1.4--2.6 mm, smooth or sometimes slightly plicate in distal
portion; stomates superficial; endostome segments 8, 1 row of cells, caducous
and inconspicuous; exostome teeth usually16, erect, papillose-granulose,
coarsely papillose or striate-papillose.
Calyptra conic-oblong,
sparsely hairy, hairs finely papillose.
Spores 10--15 \um. Usually on
non-calcareous boulders in mesic, open coniferous forests; rarely at the base
of deciduous trees; 500--3,000; Alta., B.C., Man., N.W.T., Sask., Yukon;
Alaska, Ariz., Calif., Colo., Idaho, Mont., Nev., Oreg., S.D., Utah, Wash.,
Wyo.; Europe; Asia (Japan). Orthotrichum laevigatum is a part of a complex group of
intergrading taxa that generally are characterized by an exserted, cylindric
capsule with 16 erect or reflexed exostome teeth and a poorly developed
endostome. Gametophytically, robust
plants have stiff, erect leaves and large, blunt papillae. This complex is treated here as three
species. These are O. holzingeri,
a distinct, highly specialized species of seasonally wet rocks with smooth
exostome teeth, smooth leaf cells, smooth calyptra, and lightly ribbed
capsule as distinguishing features; O.
pylaisii, distinguished by lightly
8-ribbed capsule and reflexed-recurved exostome teeth; and O. laevigatum, distinguished by smooth capsule;
erect, papillose exostome teeth. 21.
Orthotrichum lescurii
Austin, Musci Appalach., 163. 1870 Plants to 10 mm. Stem
leaves erect, stiffly appressed when dry, narrowly lanceolate to lanceolate,
2--2.5 mm, acute, margins entire, revolute to just below apex; distal laminal
cells 8--12 \um wide, 1-stratose; with 1--3 small, conic papillae; basal
laminal cells quadrate to rectangular, non-nodose. Sexual
condition goniautoicous. Seta 0.5--0.7 mm. Capsule
immersed; urn ovate-globose, constricted beneath mouth when dry, 1.1--1.8 mm,
distinctly 8-ribbed; stomates immersed in proximal half of capsule, almost
completely covered by subsidiary cells; peristome single; endostome segments
absent; exostome teeth 8, incurved-erect to spreading-recurved when dry,
densely and evenly papillose. Calyptra oblong-conic, covered with
erect, papillose hairs. Spores 12--16 \um. Calcareous
rocks, 0--1500 m; Ont.; Orthotrichum lescurii is sparsely distributed across eastern 22.
Orthotrichum lyellii
Hooker & Taylor, Musc. Brit., 76, plate 22. 1818 Dorcadion lyellii
(Hooker & Taylor) Lindberg; Orthotrichum
lyellii var. howei Renault & Cardot; O.
lyellii var. papillosum (Hampe)
Lesquereux & James; O. lyellii
var. pringlei (Müll. Hal.) Macoun
& Kindberg; O. lyellii ssp. strictum Venturi; O. lyellioides Kindberg; O.
menziesii Mitten; O. papillosum
Hampe; O. papillosum ssp. strictum (Venturi) Kindberg; O. pringlei Müll. Hal. Plants to 13 cm. Stem
leaves erect-spreading to
loosely-erect, ± contorted when dry, lanceolate to linear-lanceolate,
2.5--6.5 mm, narrowly acute to acuminate; margins plane, entire; distal
laminal cells 7--14 \um wide, 1-stratose, with conical papillae; basal
laminal cells narrowly-elongate, ± nodose. Sexual condition dioicous. Seta
0.8--2.7 mm. Capsule immersed, emergent, or rarely exserted; urn
oblong-cylindric to long-cylindric, 1.7--3 mm, strongly 8-ribbed from 1/2 to
entire length; stomates superficial; peristome double; endostome segments 16,
large, of two rows of cells, coarsely papillose; exostome teeth 16,
reflexed-recurved, densely papillose. Calyptra conic-oblong, very
hairy. Spores 18--35 \um. Usually on
trees, many times covering entire tree trunks to 10 m, rarely on boulders;
10--1000 m; B.C.; Orthotrichum lyellii is easily distinguished from all other
species of Orthotrichum by its
dioicous sexual condition, the male plants being slightly smaller and having
numerous and conspicuous perigonia. In
addition, the 16 slenderly acuminate, whitish exostome teeth and 16 endostome
segments are characteristic. Sterile
plants usually have abundant laminal brood bodies. 23.
Orthotrichum norrisii
Lara, Medina & Garilleti in Plants to 15 mm. Stem
leaves erect-appressed when dry, narrowly lanceolate to
lanceolate-ligulate from ovate base, 1.3--3 mm, narrowly obtuse, acute to
notched and channeled; margins recurved becoming incurved or plane at apex;
entire; distal laminal cells 10--16 \um wide, 1-stratose, with 2--3 simple
papillae; basal laminal cells shortly rectangular to polygonal, thin-walled
to occasionally thick-walled and nodose.
Sexual condition
goniautoicous. Seta 0.5--1 mm. Capsule emergent, urn cylindric when
dry, 1.5--2 mm, lightly 8-ribbed
the entire length; stomates immersed, in proximal third of capsule, partially
to completely covered by subsidiary cells; peristome double; endostome of 8 smooth
to faintly papillose, well-developed segments; exostome of 8 teeth,
well-developed, finely papillose, reflexed teeth. Calyptra
oblong-conic, hairy toward distal end.
Spores 14--18 \um. Epiphytic in
Mediterranean climate woodlands and chaparral; Orthotrichum norrisii is somewhat similar to O. tenellum, both with a
well-developed peristome of 8 reflexed exostome teeth and 8 long-lasting
endostome segments. Also, both species
occur on tree trunks in Mediterranean climate 24.
Orthotrichum obtusifolium
Bridel, Musc. Rec. 2(2): 23. 1801 Nyholmiella obtusifolia (Bridel) Holmen & E. Warncke; Orthotrichum inflexum Müll. Hal. Plants 3--14 mm. Stem
leaves erect-appressed when dry, ovate to ovate-lanceolate, concave, 1.4--2
mm, obtuse; margins erect-incurved, entire; distal laminal cells 13--18(--20)
\um wide, 1-stratose, unipapillose on both surfaces; basal laminal cells ±
rectangular. Sexual condition dioicous.
Seta ± 0.5 mm. Capsule
± 1/2 emergent; urn oblong-cylindric, 1.6--2 mm, strongly 8-ribbed to
base; stomates superficial at or above middle of capsule; peristome double;
endostome segments 8, composed of 2 rows of cells; exostome teeth 8,
reflexed, evenly papillose, sometimes irregularly striate above. Calyptra
short-conic, naked, papillose. Spores 20--25 \um. Deciduous
trees in open areas; in the Northeast, common on the trunks of Populus; in the West, sporadic on trunks of Juniperus, base of scrub-oaks in dry cut-over areas, and trunks
of Populus balsamifera in river bottoms; in the southern boreal forest found
commonly on aspen trunks; a pronounced
xerophyte growing only in pioneer situations; 10--2000 m; Alta., B.C., Man., N.B., Nfld., N.W.T.,
N.S., Ont., P.E.I., Sask., Que., Yukon; Alaska, Ariz., Calif., Colo., Idaho,
Iowa, Maine, Md., Mich., Minn., Mont., Neb., N.H., N.Mex. N.Y., N.C., N.Dak., Oreg., Pa.,
S.Dak., Utah, Vt., Wash., Wis., Wyo.; Europe; Asia. Species that
might be confused with Orthotrichum
obtusifolium are O. flowersii, O. exiguum, and O. gymnostomum. None of these species has distal laminal
cells with one papilla per cell as does O.
obtusifolium. In addition, O. flowersii and O. exiguum
have reflexed to revolute leaf margins and both are autoicous. Orthotrichum gymnostomum has distinctly involute leaf margins and, like O. obtusifolium,
is dioicous. 25.
Orthotrichum ohioense
Sullivant & Lesquereux in C. F. Austin, Musci Appal., 30, n. 169. 1870 Orthotrichum citrinum
Sullivant & Lesquereux; O. ohioense
var. citrinum (Sullivant & Lesquereux)
Lesquereux & James Plants 0.4--1.4 cm. Stem
leaves loosely-erect when dry, narrowly lanceolate to oblong-lanceolate,
1.4--2.9 mm; obtuse to bluntly acute; margins strongly revolute to just
before the apex, entire; distal laminal cells 8--10 \um wide, 1-stratose,
with low papillae; basal laminal cells subquadrate to short-rectangular with
rounded corners, non-nodose. Sexual condition goniautoicous. Seta
less than 1 mm. Capsule emergent; urn oblong to ovate, 1--1.5 mm, 8-ribbed to
almost base; stomates immersed; peristome double; endostome segments 8, of 1
row of cells, smooth, often not present; exostome teeth 8, later splitting to
16, reflexed, finely and densely papillose.
Calyptra oblong, naked or
slightly hairy. Spores 15--19 \um. Deciduous
trees in mesic hardwood forests especially along streams; 100--1000 m; N.B.,
N.S., Ont., Que.; Ala., Ark., Conn., Del., D.C., Fla., Ga., Ill., Ind., Ky.,
Maine, Md., Mass., Mich., Mo., Miss., N.H., N.J., N.Y., N.C., Ohio, Okla.,
Pa., S.C., Tenn., Vt., Va., W.Va. Orthotrichum ohioense
is distinguished by 16 exostome teeth, 8 rudimentary endostome segments,
lightly 8-ribbed ovate capsule, leaves that are lanceolate and bluntly acute
at apex, and cells less than 10 \um. Orthotrichum stellatum has 8 paired
exostome teeth, 8 well-developed endostome segments, and constricted,
strongly 8-ribbed capsules. 26.
Orthotrichum pallens
Bridel, Bryol. Univ. 1: 788. 1827 Orthotrichum immersum
Bridel,; O. pallens var. immersum (Bridel) Müll. Hal. Plants 3--10 mm. Stem
leaves loosely erect-appressed and slightly incurved when dry,
oblong-lanceolate to broadly lanceolate, 1.6--3 mm, narrowly obtuse, broadly
acute to broadly apiculate; margins recurved to near apex, entire; distal
laminal cells 9--14 \um wide, 1-stratose, with 2(3) low, conical papillae;
basal laminal cells rectangular, non-nodose.
Sexual condition
cladautoicous. Seta 0.5--1.5 mm. Capsule emergent to barely exserted;
urn oblong to oblong-cylindric, 1--2.5 mm, strongly 8-ribbed, becoming
narrowly-cylindric and contracted along entire length when old; stomates
immersed; peristome double; endostome segments 16, 1--2 rows of cells, smooth
or finely papillose; exostome teeth 8, erect until old, then reflexed, evenly
and densely papillose to papillae-striate.
Calyptra conic-oblong,
naked or sparsely hairy. Spores 10--20 \um. Varieties 9 (3
in the flora): North America, Orthotrichum pallens
is distinguished by having immersed stomates that are scarcely covered by thin-walled
subsidiary cells. The peristome is
double with a well-developed endostome of 16 (or 8) segments that remains
intact until the capsule is quite old.
Gametophytically, the species is distinguished by having narrowly
obtuse, oblong-lanceolate leaves, and an autoicous sexual condition. 1.
Leaf margins crenulate at the apex . . 26a. Orthotrichum pallens
var. crenulatum 1. Leaves entire. 2.
Endostome segments 8; calyptra sparsely hairy; 2.
Endostome segments 16; calyptra naked; sporadic throughout 26a.
Orthotrichum pallens var. crenulatum Vitt, Bryologist 74: 161.
1971 Stem leaves oblong to oblong-lanceolate,
rounded-obtuse, often broadly apiculate; margins usually crenulate at
apex. Seta ± 0.5 mm; urn oblong when mature, becoming strangulate when
old, 1--1.6 mm, strongly 8-ribbed nearly to base; endostome segments 8,
shorter than exostome; exostome teeth 8, reflexed, papillose-striate. Calyptra
naked. Spores 10--14 \um. Tree trunks,
especially near their base, dry, exposed areas; 500--1500 m; The endemic
variety Orthotrichum palens var. crenulatum is characterized by oblong
to oblong-lanceolate leaves with apiculate, broadly obtuse apices. The margins in almost all leaves are
crenulate near the apex and the seta is short (less than 0.6 mm). The exostome is papillose-striate as
opposed to papillose. There are 8
endostome segments and the capsule is strangulate when old. 26b.
Orthotrichum pallens
var. johnseniae Vitt, Bryologist
74: 162. 1971 Stem leaves broadly lanceolate to lanceolate,
narrowly obtuse or bluntly acute; margins entire. Seta
0.5--1 mm; urn oblong-cylindric, gradually contracted to seta through long
neck, constricted below mouth, 1.5--2 mm; endostome segments 8, of 1 row of
cells, smooth; exostome teeth 8, recurved-reflexed, finely papillose. Calyptra
sparsely hairy, the hairs smooth. Spores 13--20 \um. Basaltic boulders
in spruce-popular forests; 3000 m; Orthotrichum pallens
var. johnseniae is also endemic to
the West and is distinguished by its hairy calyptra. It has 8 endostome segments and capsules
that are shorter and moderately constricted under the mouth. The stomates are barely covered with
subsidiary cells, which usually have their interior walls thickened, and the
exostome teeth tend to be somewhat recurved instead of reflexed. 26c.
Orthotrichum pallens
Bridel var. pallens Stem leaves oblong-lanceolate to lanceolate,
narrowly obtuse to broadly acute, margins entire. Seta
0.5--1.5 mm; urn oblong-cylindric, becoming strangulate when old, 1.3--2.5
mm, endostome segments 16, long; exostome with 8, reflexed, densely
papillose. Calyptra naked. Spores 10--17 \um. Tree trunks
and branches and dry, calcareous or granitic rocks along streams; 300--3000
m; Alta., B.C., Man., Nfld., Nun., Ont., Que., Yukon; Ariz., Calif., Colo.,
Mich., Minn., Mont., Nev., S.Dak., Utah; Mexico (Nuevo León); Europe. Orthotrichum pallens var. pallens has entire leaves, 16 endostome segments, and a naked
calyptra. 27.
Orthotrichum parvulum
Mitten, J. Proc. Linn. Soc. Bot. 8: 25. 1865. Plants to 10 mm. Stem
leaves erect, stiffly appressed when dry, oblong-lanceolate to
lanceolate, 2--3.5 mm, bluntly acute to obtuse; margins entire, plane or
weakly recurved; distal laminal cells 10--15 \um wide, 1-stratose; with 1--2
low, conic papillae; basal laminal cells quadrate to rectangular, non-nodose. Sexual
condition goniautoicous. Seta 0.5--0.8 mm. Capsule
immersed; urn oblong, constricted beneath mouth when dry, 1.2--1.9 mm,
distinctly 8-ribbed; stomates immersed in distal half, almost completely
covered by subsidiary cells; peristome single or rarely with fugacous endostome;
endostome segments 8 when present, thin, short, hyaline; exostome teeth 16,
incurved-erect to spreading-recurved when dry, coarsely papillose to
reticulate-papillose. Calyptra oblong-conic, covered with
erect papillose hairs. Spores 12--19 \um. Calcareous
rocks; 90--130 m; Orthotrichum parvulum is most frequent in the interior
highlands of 28.
Orthotrichum pellucidum
Lindberg, Oefv. K. Svensk. Vet. Ak. Foerh. 23: 549. 1867 Orthotrichum cupulatum
var. jamesianum (James) Plants to 2.5 cm. Stem
leaves stiff, closely erect-appressed, incurved when dry,
oblong-lanceolate or lanceolate, 2--2.7 mm, blunt or obtuse; margins recurved
below, reflexed in distal portion, entire; distal laminal cells 6--11 \um
wide, 1-stratose, papillae always large, conical or forked; basal laminal
cells short-rectangular to quadrate, grading to subquadrate at margin. Sexual
condition goniautoicous. Seta 1--1.5 mm. Capsule
1/2 emergent to shortly exserted;
urn oblong to short-oblong when old and dry, oblong-ovate to ovate when
mature, 1.2--1.9 mm, strongly 8-ribbed entire length, rarely 16-ribbed with 8
short intermediate ribs; stomates immersed in middle and proximal portion, ±
completely covered by protruding subsidiary cells; peristome single;
endostome segments none; exostome teeth usually 16 when mature, irregularly
split, often absent when capsule is old, papillose-striate to coarsely
reticulate-papillose. Calyptra oblong, covered with
papillose, whitish hairs. Spores 10--15 \um. Calcareous,
more rarely siliceous boulders and cliff faces in xeric areas, often found
growing in direct sunlight; 500--3000 m; Greenland; Alta., B.C., Man.,
N.W.T., Nun., Yukon; Alaska, Ariz., Calif., Colo., Idaho, Mont., Nev., N.Mex.,
Oreg., S.Dak., Utah, Wyo.; n Europe. Orthotrichum pellucidum
is distinguished by the blunt, lanceolate-oblong leaves that always have a
glaucous appearance because of the large forked papillae of the distal
laminal cells. When the plants are
fertile, which is usually the case, the shortly exserted, oblong-ovate,
eight-ribbed capsule should identify this species. 29.
Orthotrichum praemorsum
Venturi in J. Roell, Bot. Centralbl. 44: 418.
1890 Orthotrichum affine
ssp. praemorsum (Venturi) Kindberg;
O. arcticum var. praemorsum (Venturi) Wijk &
Margadant; O. microblephare var. praemorsum (Venturi) Grout Plants 1--2.5 cm. Stem
leaves erect-appressed when dry,
ovate-lanceolate to narrowly lanceolate, 2.5--4.5 mm, gradually narrowed to acuminate-cuspidate
apex; margins recurved-revolute to near apex, entire; distal laminal cells
9--13 \um wide, 1-stratose, with 1--3 low, forked papillae; basal laminal
cells elongate-rectangular, nodose. Sexual condition goniautoicous. Seta
0.5--1.1 mm. Capsule 1/2 emergent when dry, barely emergent when moist; urn
ovate-oblong when mature, oblong to oblong-cylindric and constricted beneath
mouth when old and dry, 1.6--2.4 mm, strongly 8-ribbed 1/2 to 2/3 the length;
stomates superficial; peristome double; endostome segments 16, large,
coarsely papillose; exostome teeth 8, reflexed-recurved, finely papillose,
sometimes papillose-striate below. Calyptra short-conic, hairy, hairs
with small papillae. Spores 12--18 \um. Rocks, rarely
lava, dry montane areas; middle elevations; Orthotrichum praemorsum is rare, having been collected only a
few times. It is distinguished from other species by strongly-ribbed,
emergent capsule, and by the cuspidate, shortly hyaline-tipped leaves. 30.
Orthotrichum pulchellum
Brunton in J. E. Smith, Engl. Bot. 25: (Sept. 1) 1807 Orthotrichum pulchellum
var. groutii Plants to 1.6 cm. Stem
leaves loosely-twisted and crisped-flexuose when dry, narrowly lanceolate
to linear-lanceolate, 1.6--3.2 mm, sharply acute; margins recurved to near
apex, entire; distal laminal cells 8--14 \um wide, 1-stratose, with small,
conical papillae; basal laminal cells rectangular, non-nodose. Sexual
condition goniautoicous. Seta (0.8--)1--3(--3.5) mm. Capsule
exserted or sometimes 1/2 emergent; urn oblong-cylindric, 0.9--2.1 mm,
strongly 8-ribbed entire length, stomates immersed in middle and distal
portion of capsule, 1/2 to completely covered by strongly differentiated
subsidiary cells; peristome double; endostome segments 16, of 1 row of cells,
finely roughened; exostome teeth 16, rarely united into 8 pairs, erect when
mature, later reflexed, densely papillose.
Calyptra oblong-conic to
conic, naked. Spores 12--24 \um. Tree trunks,
branches, and twigs of deciduous and coniferous trees; 0--1000 m: B.C.; Orthotrichum pulchellum is distinguished from all species of Ulota by glabrous calyptra and
immersed stomates, and from O. consimile by 16, reddish exostome
teeth that are papillose-reticulate with slender, high papillae, and by
stomates that are always located at or beyond the middle of the capsule.
Also, O. pulchellum plants are rarely longer than 1 cm and capsules are
usually just barely exserted and oblong-cylindric. 31.
Orthotrichum pumilum Swartz, Monthly Rev. London 34:
538. (June 1) 1801 Orthotrichum affine
var. fallax (Bridel) Hampe; O. brachytrichum Lesquereux &
James; O. fallax Bridel; O. fallax var. truncatulum Austin; O.
pumilum var. americanum
Venturi; O. pumilum var. fallax (Bridel) J. Kickx f.; O. pumilum var. fallax (Bridel) Bertsch; O.
schimperi Hammar; O. schimperi
var. truncatulum (Austin) Paris; O. tenellum var. pumilum (Swartz) Boulin Plants to 5 mm. Stem
leaves loosely-erect when dry, ovate-lanceolate to lanceolate, 1.8--2.8
mm, sharply to slenderly acute, most leaves ending in 1--2 celled apiculus,
rarely obtuse; margins revolute nearly to apex, entire; distal laminal cells
14--20 \um wide, 1-stratose, smooth or with 1--2 small, conical papillae;
basal laminal cells rectangular, grading to quadrate at margin,
non-nodose. Sexual condition autoicous.
Seta less than 0.5 mm. Capsule
immersed to 3/4 emergent; urn oblong to oblong-cylindric, 1.2--1.5 mm,
strongly 8-ribbed; stomates immersed, at middle of capsule, 1/2 or less
covered by subsidiary cells; peristome double; endostome segments 8, of 1--2
rows of cells, smooth; exostome teeth 8, rarely splitting to 16, reflexed,
finely and evenly papillose. Calyptra oblong, sparsely hairy or
rarely naked. Spores 14--21 \um. Trunks, lower
branches, and bases of deciduous trees, rarely on coniferous trees or in
crevices of rocks; common in open, hardwood forests and on trees along
roadsides; 10--1000 m; Alta., Man., Ont., Sask.; Ariz., Calif., Colo., D.C.,
Idaho, Ill., Iowa, Kans., Md., Mass., Mich., Minn., Mis., Mo., Mont., Neb.,
N.H., N.J., N.Mex., N.Y., N.Dak., Ohio, Okla., Pa., S.C., S.Dak., Tenn.,
Texas, Utah, Vt., Va., Wash., W.Va., Wisc.; Europe; n Africa. Orthotrichum pumilum
can usually be distinguished by its strongly ribbed capsules, reflexed
peristome teeth, apiculate leaves with cells larger than 14 \um, and papillae
that are very small or absent. In
eastern 32.
Orthotrichum pusillum
Mitten, J. Linn. Soc. Bot. 8: 25. (June 30) 1864 Orthotrichum psilocarpum
Sullivant Plants 2--4 mm. Stem
leaves loosely-appressed when dry, ovate-lanceolate to oblong-lanceolate,
1--2.2 mm, narrowly obtuse to broadly acute, sometimes with small apiculus;
margins revolute nearly to apex, entire except for denticulate apex on some
leaves; distal laminal cells 9--14 \um wide, 1-stratose, with 2--4 small,
conical papillae; basal laminal cells, broadly rectangular, non-nodose. Sexual
condition autoicous. Seta less than 1 mm. Capsule
immersed to 1/2 emergent; urn oblong to ovate-oblong, 0.8--1.4 mm,
wrinkled to lightly 8-plicate; stomates immersed, at middle or distal portion
of capsule; peristome single; endostome segments usually absent or if present
8, rudimentary, smooth; exostome teeth 16, erect or recurved. Calyptra
oblong, smooth. Spores 13--20 \um. Trunks of
deciduous trees in open, dry woods and along roadsides, rarely on Juniperus associated with habitation
along streams; 10--1000 m; Ala., Ark.,
Conn., D.C., Fla., Ga., Ill., Ind., Kans., Ky., La., Md., Mass., Mich.,
Miss., Mo., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., S.C. Tenn., Texas, Va.,
W.Va. Orthotrichum pusillum
is distinguished by its smooth, oblong capsule, which is sharply contracted
to the seta, 16 peristome teeth, which are erect or recurved when old, some
leaves of each plant having 1--4 small denticulations at the apex, and distal
laminal cells 9--14 \um in width. The
leaf cells are smaller than those of O.
pumilum and in longitudinal rows,
but they are larger, with thinner walls, than those of O. ohioense. To help identify sterile material: O. pusillum
has denticulate leaf apex that may be narrowly obtuse or acute, and leaf cells 9--14 \um with thin walls; O.
pumilum has an apiculate leaf apex
that is normally acute and entire, and leaf cells 14--20 \um, with thin
walls; and O. ohioense has
an obtuse leaf apex, which is never apiculate or denticulate, and leaf cells
8--10 \um, with thick walls. 33.
Orthotrichum pycnophyllum
Müll. Hal., Syn. 1: 709. 1849 Orthotrichum lozanoi
Cardot; O. lozanoi var. lutescens Cardot; O. pycnophyllum var. lutescens
(Cardot) Grout; O. recurvans Müll.
Hal. Plants 0.8--4 cm. Stem
leaves loosely erect-spreading when
dry, lanceolate, 2.5--4 mm, gradually acuminate to long-acuminate-cuspidate;
margins revolute to just below apex, often wavy and slightly notched near
apex; distal laminal cells 7--13 \um wide, 1-stratose, with 1--2 conical
papillae per cell; basal laminal cells elongate, nodose. Sexual
condition goniautoicous. Seta 1--1.5 mm. Capsule
immersed to fully exserted, generally 1/2--1/4 emergent, urn oblong-ovate to
cylindric, 1.5--2 mm, smooth to lightly 8-ribbed; stomates superficial;
peristome double; endostome segments 16, 2 cells wide, coarsely papillose,
usually not erose; exostome teeth 16, recurved, obscurely and densely
papillose. Calyptra conic-oblong, moderately hairy, with dark beak. Spores
17--29 \um. Branches and
trunks of trees; 500--2000 m; Orthotrichum pycnophyllum is restricted to southwestern North
America, being found only south of 35° N latitude in the southwestern 34.
Orthotrichum pylaisii
Bridel, Brol. Univ. 1: 722. 1826 Orthotrichum arcticum
Schimper; O. arcticum ssp. blyttii (Schimper) Kindberg; O. arcticum var. groenlandicum (Berggren) Moenkemeyer; O. blyttii Schimper; O.
blyttii ssp. arcticum
(Schimper) Macoun; O. blyttii ssp. microblephare (Schimper) Macoun; O. blyttii var. arcticum (Schimper) I. Hagen; O.
blyttii var. groenlandicum
(Berggren) C. E. O. Jensen; O. blyttii
var. microblephare (Schimper) C. E.
O. Jensen; O. groenlandicum
Berggren; O. idaohense Cardot &
Thériot; O. pylaisii ssp. arcticum (Schimper) Kindberg; O. pylaisii ssp. groenlandicum (Berggren) Kindberg; O. pylaisii ssp. microblephare
(Schimper) Kindberg; O. rhabdophorum
Venturi Plants to 3 cm. Stem
leaves erect-appressed when dry,
ovate-lanceolate to lanceolate, 2--3 mm, broadly acute to narrowly obtuse, often
bluntly acuminate; margins revolute, entire; distal laminal cells 9--16 \um
wide, 1-stratose, with 1--4 forked papillae; basal laminal cells
rectangular-elongate, ± nodose, thick-walled.
Sexual condition goniautoicous. Seta
2--3 mm. Capsule emergent or exserted; urn oblong to cylindric, 1.2--2.5
mm, moderately 8-ribbed ± 1/2 length; stomates superficial; peristome double
or single; endostome segments 8, about 1/2 as long as exostome, narrow, of 1
row of cells, smooth or finely papillose, often absent; exostome teeth 16,
sometimes united into 8 pairs, 4--6 cells wide, densely papillose. Calyptra
oblong to oblong-conic, sparsely hairy.
Spores 16--27 \um. Usually on
rocks near the ocean or in alpine or arctic zones, rarely on trees such as Salix or Alnus, bone and frequently in or near bird rookeries, in western
North America found on granitic boulders in coniferous forests; 100--3000 m;
Greenland; B.C., Alta., Man., Nfld., N.W.T. Nun., Que., Yukon; Alaska,
Calif., Idaho, Mont., Wash., Wyo.; n Europe; n Asia. Orthotrichum pylaisii is distinguished by ribbed capsules with
reflexed exostome teeth and a rudimentary endostome, along with superficial
stomates and large, ovate-lanceolate leaves.
The capsule is almost always exserted in North America; in 35.
Orthotrichum rivulare
Turner, Musc. Hib., 96. 1804 Plants 1--5 cm; in loose, dark green to
blackish tufts. Stem leaves erect and loosely appressed when dry;
oblong-lingulate, oblong-lanceolate, or ovate-lanceolate, 2.8--5 mm, broadly
rounded to obtuse, flat; margins strongly revolute to just below apex,
usually irregularly dentate at apex; distal laminal cells 10--13 \um wide,
1-stratose, with 1--3 small, conical papillae; basal laminal cells rectangular
to short-rectangular, smooth. Sexual condition goniautoicous. Seta
1--1.5 mm. Capsule immersed when moist, barely to 3/4 emergent when dry; urn
oblong to oblong-ovate, 1.5--2.5 mm, strongly 8-ribbed entire length;
stomates immersed; peristome double; endostome segments 16, usually 1 cell
wide, finely papillose; exostome teeth 8, sometimes splitting to 16, erect
until old, then loosely reflexed, irregularly papillose, papillae tending to
be confluent forming longitudinal striae.
Calyptra 16-plicate with
long beak, naked. Spores 10--17 \um. Exposed tree roots, bases of trees along streams, also commonly on siliceous boulders at the edges of streams and rivers, occasionally on rocks that are frequently inundated, rarely in aquatic habitats; 0--1000 m; Alta., B.C.; Calif., Idaho, Nev., Oreg., Wash., Wyo.; Europe. Orthotrichum rivulare is distinguished from all others in
the genus by the following combination of characters: (1) leaves often obtuse, the margins
dentate near the apex, (2) calyptra plicate, the projecting edges papillose,
otherwise smooth and naked, (3) 16 endostome teeth, and (4) stomates
completely covered by the overarching subsidiary cells. From O.
euryphyllum, the flat, dentate apex is a key differentiating feature. 36.
Orthotrichum rupestre
Schwaegrichen, Spec. Musc. Frond. Suppl. 1(2): 27. 53. 1816 Dorcadion rupestre
(Schwaegrichen) Lindberg; Orthotrichum
bullatum Müll. Hal.; O. californicum Venturi; O. douglasii Duby; O. macfaddenae R. S. Williams; O. rupestre var. globosum (Lesquereux) Grout; O.
rupestre var. macfaddenae (R.
S. Williams) Grout; O. texanum Sullivant & Lesquereux; O. texanum var. globosum Lesquereux Plants 1--12.5 cm, usually longer than 3
cm. Stem leaves erect-appressed when dry, narrowly lanceolate to
ovate-lanceolate, 2--4.5 mm, sharply to slenderly acute; margins recurved to
narrowly revolute to near apex, entire; distal laminal cells 6--13 \um wide,
1-stratose, with 1--2 small, conical or low, forked papillae per cell; basal
laminal cells elongate, thick-walled and nodose to rectangular and
thin-walled. Sexual condition
goniautoicous. Seta less than 1.8 mm. Capsule
immersed, emergent or slightly exserted, usually ± 1/3 emergent; urn globose
ovate, ovate-oblong, or short-oblong, 1.3--1.8 mm, lightly or distinctly
8-ribbed for 1/2--2/3 the length; stomates superficial; peristome single,
rarely double; endostome segments lacking or rarely 8 and rudimentary; smooth
or roughened; exostome teeth 16, erect or sometimes spreading-recurved when
old; smooth, scattered-papillose to coarsely papillose-granulate. Calyptra
oblong, hairy or rarely naked, hairs finely papillose. Spores
13--21 \um. Non-calcareous boulders and cliff faces in mesic areas of pine, spruce-fir or aspen forests, usually at higher elevations, rarely found in sterile condition at the base of trees, frequently collected in subalpine shaded situations; 100--3000 m; Greenland; Alta., B.C., N.W.T., Nun., Ont., Yukon; Alaska, Ariz., Calif., Colo., Idaho, Mont., Nev., N.Mex., Oreg., S.Dak., Utah, Wash., Wyo.; South America ; Europe; Asia; n and e Africa; Asia (Japan); Atlantic Islands (Canary Islands); Pacific Islands (New Zealand); Australia. In typical
form, O. rupestre is characterized as follows: plants much branched in
loose tufts, leaves lanceolate, apex narrowly and bluntly acute, basal cells
elongate, thick-walled, and nodose, yellowish in older leaves, capsule
oblong, emergent, 8-ribbed 1/2 the length when old and dry, smooth or
slightly 8-ribbed when mature, exostome of 16 erect teeth, coarsely
papillose, and calyptra with abundant spinulose, papillose hairs, some of
which extend over the beak. 37.
Orthotrichum shawii
Plants to 4 cm. Stem
leaves erect-appressed when dry, lanceolate, 2.7--3.6 mm, shortly
acuminate to acute, margins broadly recurved, plane at apex, smooth; distal
laminal cells 9--15 um wide, 1-stratose, with 2--4 thick, well-developed,
blunt, simple or forked papillae; basal laminal cells long-rectangular, with
thick and nodose walls. Sexual condition goniautoicous. Seta
less than 0.4 mm. Capsule immersed to emergent; urn ovate to cylindric-ovate,
1.6-1.8 mm, usually smooth or lightly 8-ribbed when dry; stomates superficial
in middle and distal portion of capsule, peristome usually single; endostome
rarely present, of 8 very short hyaline segments; exostome teeth 16,
irregularly divided and recurved, coarsely papillose to
papillose-striate. Calyptra oblong-conic, with long,
yellowish, papillose hairs. Spores 14--17(--20) mm. Apparently on
tree trunks; Orthotrichum shawii is similar to O. rupestre, differing in an exostome of 16 short, recurved teeth
and nearly smooth capsules compared to O.
rupestre with erect or spreading exostome teeth and variable capsule
ribbing. This species was collected once without locality, by Bolander in the
1860’s, but is probably not uncommon in 38.
Orthotrichum shevockii
Lewinsky-Haapasaari & D. H. Norris, Bryologist 101: 435. 1998 Plants less than 1.5 cm. Stem
leaves rigid, erect to spreading
when moist, ligulate- to elliptic-lanceolate, less than 5. mm, obtuse to
rounded-acute; margins recurved from near apex to just above base, entire;
distal laminal cells 13--16 \um wide, 1-stratose, strongly papillose,
papillae high, simple to forked; basal laminal cells short-rectangular,
pellucid and thin-walled with corner thickenings. Sexual
condition goniautoicous. Seta less than 0.5 mm. Capsule
immersed; urn short-cylindric to oblong-ovate, 1--2 mm, strongly 8-ribbed
entire length; stomates immersed, 2/3 to completely covered by well-developed
subsidiary cells; peristome double; endostome segments 8 well-developed,
occasionally 16, 1 cell wide, smooth or roughened; exostome teeth 8 ± 16,
reflexed, papillose to papillose-reticulate near apex. Calyptra
oblong-conic, hairy, hairs papillose. Spores 15--18 \um. Dry granitic
rocks; 1100--1600 m; Orthotrichum shevockii is known only from the southern 39.
Orthotrichum sordidum
Sullivant & Lesquereux in C. F. Austin, Musci Appal., 30. n. 168. 1870 Orthotrichum affine
var. sordidum (Sullivant &
Lesquereux ) Grout; O. affine ssp. subrivale Kindberg; O. cancellatum Cardot & Thériot Plants 0.5--3 cm. Stem
leaves loosely erect-appressed and
often slightly contorted when dry, narrowly lanceolate to oblong-lanceolate,
2--3.6 mm, sharply acute or bluntly acuminate; margins revolute to near apex,
entire; distal laminal cells 7--14 \um wide, 1-stratose, with 2--3 conical
papillae per cell; basal laminal cells elongate-rectangular, nodose,
thin-walled. Sexual condition goniautoicous.
Seta 0.5--1.6 mm. Capsule
usually 1/2--2/3 emergent, sometimes immersed or fully exserted, tapering
to seta through a long, shriveled neck, which is less than 1.5 mm in length;
urn oblong to oblong-cylindric, 0.8--1.5 mm, strongly 8-ribbed 1/2--2/3
length; stomates superficial; peristome double; endostome segments 8, 1--2
rows of cells, smooth or finely verrucose; exostome teeth 8, reflexed,
reticulately papillose to obscurely papillose. Calyptra
oblong-conic, sparsely hairy, hairs smooth or with small papillae. Spores
19--29 \um. In the Great
Lakes region usually trunks of both deciduous and coniferous trees, rarely
boulders in hardwood forests, in the Arctic usually on small trees; 0--1000
m; Man., N.B., Nfld., N.W.T., N.S., Ont., P.E.I., Que.; Alaska, Maine, Mass.,
Mich., Minn., N.H., N.J., N.Y., Pa., Vt., Wis.; n Asia. Orthotrichum sordidum species is distinguished by having oblong capsules with exostome teeth perforate almost to the base and cancellate in the distal portions. 40.
Orthotrichum speciosum
Nees in J. W. Sturm, Deutschl. fl. 2(3) (fasc. 17): 5. 1819 Orthotrichum killiasii
Müll. Hal.; O. speciosum ssp. killiasii (Müll. Hal.) Kindberg; O. speciosum var. killiasii (Müll. Hal.) Venturi Plants 0.8--5 cm, robust. Stem
leaves loosely erect, usually slightly contorted or curved, lanceolate to
narrowly lanceolate, 3--4 mm, acuminate to narrowly acute, margins revolute
nearly to apex, entire; distal laminal cells 7--13 \um wide, 1-stratose,
1--3, conic or forked papillae; basal laminal cells elongate, with nodose or
wavy walls. Sexual condition goniautoicous.
Seta 1--3 mm. Capsule
exserted to rarely emergent; urn cylindric to ovate-cylindric when mature,
1.5--2.4 mm, lightly ribbed ± 1/2 length of capsule; stomates superficial; peristome
double; endostome segments 8, 2 rows of cells, coarsely papillose; exostome
teeth 8, recurved, coarsely papillose.
Calyptra conic-oblong, with
finely papillose hairs. Spores 13--26 \um. Coniferous and decidious
trees, rocks; 10--2000 m; Greenland; Alta., B.C., Man., N.B., Nfld., N.W.T.,
N.S., Nun., P.E.I., Que., Sask. Yukon; Alaska, Calif., Colo., Idaho, Mich.,
Minn., Mont., Nev., N.H., N.Y., Oreg., Utah, Wash., Wyo.; Europe, n Africa; n
Asia. Orthotrichum speciosum
is mainly western and Arctic in distribution.
The ribbed capsule, large size, and more slender leaf apices
distinguish it. Orthotrichum elegans has smooth
capsules. 41.
Orthotrichum spjutii
Norris &Vitt, Nova Hedw. 56: 259. 1993
Plants to 3.5 cm. Stem
leaves erect-spreading to
spreading when moist, loosely erect-appressed to erect-curved when dry, 2--4
mm, broadly lanceolate, ovate-lanceolate, or ovate-ligulate, bluntly acute to
obtuse; margins strongly revolute from just before apex to near base,
papillose-crenulate; distal laminal cells 10--15 \um wide, 1-stratose,
irregularly rounded to isodiametric, with obvious corner thickenings,
strongly papillose, papillae forked and densely arranged; basal laminal cells
long-rectangular to elongate, pellucid, smooth, lateral walls somewhat
irregularly thickened, shorter, narrower, somewhat papillose along the
margins. Sexual condition goniautoicous.
Seta 1--1.5 mm. Capsule
emergent, often rather shortly so; urn oblong to shortly-cylindric,
erect, nearly smooth to lightly 8-ribbed sometimes becoming deeply 8-ribbed
and constricted with flaring mouth when old and dry; stomates superficial;
peristome superficially absent, of about 16, very short, blunt, papillose
teeth 1--3 cells high. Calyptra ovoid-campanulate, mitrate,
plicate, hairy; hairs with spiculose papillae throughout. Spores
8--11 \um, finely papillose. Shaded forests near stream
beds, canyons, usually in mountainous areas, deciduous trees, rarely on
shaded rocks; 2600 m; Orthotrichum spjutii
is confined to California, and is best recognized by the following
combination of characters: large
multifid distal leaf cell papillae, strongly revolute leaf margins, emergent,
oblong capsules with peristome reduced to 1--2 cells and seemingly absent,
and small spores. 42.
Orthotrichum stellatum Bridel, Bryol. Univ. 1: 274. 1826 Orthotrichum braunii Bruch, Schimper & W. Gümbel Plants 3--8 mm. Stem
leaves loosely-erect when dry, lanceolate to oblong-lanceolate, 1.8--2.2 mm,
narrowly obtuse to bluntly acute; margins recurved to just below the apex,
entire; distal laminal cells 8--13 \um wide, smooth or with small, conic
papillae; basal laminal cells rectangular, becoming nodose toward the costa. Sexual
condition goniautoicous. Seta 0.4--1 mm. Capsules
emergent; urn oblong-cylindric and strangulate when dry, 0.4--1 mm,
dark-brown ribs; stomates immersed; peristome double; endostome segments 8,
of 1--2 rows of cells, smooth; exostome teeth 8, reflexed finely and evenly
papillose. Calyptra oblong, naked or sparsely hairy. Spores
13--18 \um. Hardwood tree
trunks in moist forests, often associated with O. ohioense in mesic, shaded situations along streams; 10--1000
m; Nfld, N.B., N.S. Ont.; Ark., Del., Ga., Ill., Ind., Iowa, Ky., Maine, Md.,
Mass., Mich., Mo, N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C.,
Tenn., Ver., Va., W.Va. Orthotrichum stellatum is distinguished by strongly ribbed,
strangulate capsules with a well-developed endostome of 8 segments. Orthotrichum
ohioense has weakly ribbed capsules that are not constricted beneath the
mouth and an endostome which is lacking or rudimentary. From O.
pumilum, the narrowly lanceolate leaves and small leaf cells with
unevenly incrassate walls differentiate O.
stellatum. 43.
Orthotrichum stramineum Bridel, Bryol. Univ., 789.
1827 Orthotrichum strangulatum var. stellatum (Bridel) E. Britton Plants 0.5--1 cm. Stem
leaves erect-appressed or slightly flexuose when dry, lanceolate,
1.8--2.4 mm, acute; margin recurved to just below apex, entire; distal
laminal cells 8--12 \um wide, 1-stratose, smooth or small, conic papillae;
basal laminal cells rectangular, smooth, rather thin. Sexual
condition goniautoicous. Seta 0.7--1.5 mm. Capsules
emergent, elliptic-cylindric and strangulate, urn 0.8--1.5 mm, strongly
8-ribbed; stomates immersed, 1/2 to completely covered by well-developed
subsidiary cells; peristome double, endostome segments 8 or 16, 1--2 rows of
cells wide, smooth on outside or finely sriate; exostome teeth 8, reflexed to
revolute when old and dry, finely and evenly papillose. Calyptra
oblong, with few smooth or papillose hairs, rarely naked. Spores
10--15 \um. Trees; 10--500 m; Nfld.;
Europe; Orthotrichum stramineum is distinguished by rather long,
elliptic (fat), strongly ribbed capsules that are strangulate as useful field
characters. The hairy vaginula and the
shape of the capsule will separate O.
stramineum from O. stellatum. 44.
Orthotrichum strangulatum Palisot de Beauvois, Prodr., 81.
1805 Orthotrichum calcareum Austin; O. lescurii ssp. porteri
(Sullivant & Lesquereux) Kindberg; O.
missouricum Grout; O. nudum
var. strangulatum (Palisot de
Beauvois) Baehni; O. peckii
Sullivant & Lesquereux; O. porteri
Sullivant & Lesquereux; O.
strangulatum var. typicum
Venturi; O. strangulatum var. missouricum Grout Plants 4--10 mm. Stem
leaves erect, stiff when dry, ovate- to oblong-lanceolate, 2--2.7 mm,
acute to ± narrowly obtuse; margins thickened to near apex, entire; distal
laminal cells 7--12 \um wide, 1-stratose (or occasionally with 2-stratose
streaks) except at margins, there 2- to 3(--7)-stratose, with 2--3 conic,
sometimes forked papillae; basal laminal cells quadrate to rectangular,
non-nodose. Sexual condition goniautoicous.
Seta 0.8 mm or less in
length. Capsules immersed to 1/2
emergent; urn oblong-ovate to oblong, 1.1--1.8 mm, distinctly 8-ribbed and
constricted beneath mouth when dry; stomates immersed in distal half of
capsule, nearly completely covered by subsidiary cells; peristome single (or
rarely double); endostome segments none (or rarely 8 fragile, short
segments); exostome teeth 16, incurved-erect to spreading-recurved, granulose
or reticulate-papillose. Calyptra oblong-conic, covered with
erect papillose hairs. Spores 12--16 \um. Found
exclusively on dry, exposed, calcareous or dolomitic bluffs and rock faces;
150-700 m elev; Man., Ont., Que.; Ala., Ark., Ill., Ind., Iowa, Kans., Ky.,
Maine, Md., Minn., Mo., N.J., N.Y., N. C., Ohio, Okla., Pa., S.C., S.D.,
Tenn., Vt., Va., W.Va. The Orthotrichum strangulatum complex
contains three species, all endemic to eastern and middle 45.
Orthotrichum striatum
Hedwig, Spec. Musc. Frond., 163. 1801 Dorcadion striatum
(Hedwig) Lindberg; Weissia striata (Hedwig) P. Gaertner, B. Meyer
& Schreber Plants 1--6 cm, robust. Stem
leaves loosely-erect, flexuose when dry, lanceolate, 3--4 mm, narrowly
acute to acuminate, appearing revolute to near apex, margins entire; distal
laminal cells 7--9 \um wide, 1-stratose, with simple papillae; basal laminal
cells elongate, nodose. Sexual condition goniautoicous. Seta
0.8--1.2 mm. Capsule immersed; urn broadly-oblong to narrowly-ovate, 1.5--2.3
mm, smooth; stomates superficial; peristome double; endostome segments 16, in
2 rows of cells, densely papillose; exostome teeth 16, recurved, densely
papillose. Calyptra broadly oblong, moderately hairy. Spores
17--31 \um. Deciduous
trees, usually Acer and Alnus trunks in moist woods, less
commonly on conifers; 10--1000 m; B.C.; Alaska, Calif., Idaho, Wash., Oreg;
Europe; n Africa Asia (India in Kashmir). Orthotrichum striatum is distinguished from its closest
relatives, O. pycnophyllum, O. speciosum, and O. lyellii, by its
ovate, smooth capsule, which is immersed even when dry. The exostome consists of 16, recurved
teeth, and the endostome consists of 16 erose segments. In the area of the flora, Orthotrichum striatum is restricted to the 46.
Orthotrichum tenellum
Bridel, Bryol. Univ. 1: 786. 1827 Orthotrichum coulteri
Mitten; O. cylindrocarpum
Lesquereux; O. tenellum var. coulteri (Mitten) Grout; O. tenellum var. cylindrocarpum (Lesquereux) L. F. Koch Plants 3--10 mm. Stem
leaves stiffly erect-appressed, tips incurved when dry, spreading to wide
spreading in distal portions of branches, ranging from ligulate to
lanceolate-oblong, 1.3--2.6 mm, polymorphic, obtuse to broadly acute, rarely
ending in short, hyaline cell, apiculate, or serrulate, concave; margins
revolute to below apex or sometimes broadly reflexed in distal 1/2 of leaf,
entire; distal laminal cells 8--13 \um wide, 1-stratose, with 2--3 conical
papillae per cell; basal laminal cells broadly rectangular, non-nodose. Sexual
condition usually cladautoicous, rarely goniautoicous. Seta
less than 2 mm. Capsule 2/3 emergent to shortly-exserted; urn cylindric to
cylindric-elliptic when mature, 1.3--2 mm, strongly 8-ribbed entire length;
stomates immersed in proximal 1/3 of capsule, ± completely covered by
projecting subsidiary cells; peristome double; endostome segments 8, rarely
16, finely papillose, exostome teeth 8, rarely split to 16 when old, erect
when mature, reflexed when old. Calyptra conic-oblong, naked or
sparsely hairy, the hairs sometimes papillose by projecting walls. Spores
11--18 \um. Trunks of
deciduous trees in dry, open areas, rarely on rocks; restricted to lower
elevations; 100--2000 m; B.C.; Calif., Nev., Wash.; Europe, n Africa;
Atlantic Islands (Azores, Canary Islands). Orthotrichum tenellum
is the only western species with deeply immersed stomates, blunt leaves, and
oblong-cylindric capsules. Its small
size distinguishes it from other Californian species found on trees. 47.
Orthotrichum underwoodii F. Lara, R. Garilleti & V. Mazimpaka,
Bryologist 104: 268. 2001 Plants to 1 cm. Stem
leaves erect when dry, slightly flexuose to erect when moist, lanceolate,
2--3.2(--3.5) mm, with acute apex; margins entire or papillose-crenulate;
distal laminal cells (7--)11--14(--16) \um wide; 1-stratose, basal laminal
cells rectangular to almost quadrate, rarely nodose. Sexual
condition goniautoicous or cladautoicous.
Seta 0.4--0.6 mm. Capsule
2/3 emergent to shortly-exserted; urn cylindric, 8 narrow ribs entire length;
stomates immersed; peristome double; endostome segments 16, filiforme,
exostome teeth 16, erect when mature, reflexed when old. Calyptra
conic-oblong, slightly plicate, with scattered short, thin hairs. Spores
16--18(--19) \um. Trees;
elevation unknown; Orthotrichum
underwoodii is known only
from type material from the For Key, see
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1.
Leaves gradually acute and ending in hyaline awn .
. 9. Orthotrichum diaphanum
1. Leaves obtuse,
acute, acuminate, or cuspidate, not ending in hyaline awn.
2. Leaf margins erect-incurved, or involute; leaves
ovate or elliptic, concave with broadly obtuse or rounded apices; laminal
brood-bodies abundant; dioicous; calyptrae papillose, neither plicate nor
hairy.
3. Leaf margins involute; distal laminal cells
with 2--3 conical papillae per cell; peristome lacking;
3. Leaf margins erect-incurved to incurved;
distal laminal cells with 1 papilla per cell; peristome double; common
throughout boreal and western regions . .24. Orthotrichum obtusifolium
2. Leaf margins plane, recurved, revolute, or
thickened; leaves ± lanceolate or oblong, plane to keeled, with narrowly
obtuse, acute, or acuminate apices; laminal brood bodies absent, few, or
abundant; autoicous, if dioicous then leaves acuminate; calyptrae smooth,
rarely papillose, plicate, hairy or naked.
4. Leaves crisped-flexuose when dry; capsule
cylindric or elliptic-cylindric, fully exserted; stomates immersed; Pacific
Northwest and
5. Plants 8--20 cm; distal laminal cells 7--11 \um
wide; seta usually 2.5--6 mm; capsule long-exserted, elliptic-cylindric when
mature; peristome teeth 8, light-brown, finely papillose on the exterior,
finely striate on the interior; stomates in the middle and proximal portion of
the capsule . . 7. Orthotrichum consimile
5. Plants usually less than 1.3 cm; distal
laminal cells 9--14 \um wide; seta 1.5--3.5 mm; capsule barely exserted,
oblong-cylindric when mature; peristome teeth 16, red, irregularly
papillose-reticulate; stomates above the middle of the capsule, never in the
proximal portion . . 30. Orthotrichum
pulchellum
4. Leaves erect-appressed when dry; capsules
various; stomates superficial or immersed; throughout
6. Leaves with abundant laminal gemmae;
restricted to
6. Leaves without or with few laminal gemmae;
variously distributed in
7. Stomates superficial; basal leaf cells
elongate, ± nodose, thick-walled.
8. Endostome segments well-developed, usually present
when capsule is old; exostome teeth usually recurved or reflexed; prostome
absent; leaves sometimes flexuose when dry, 1-stratose
9. Exostome teeth reflexed; endostome segments 8,
narrow, consisting of a single row of cells; capsule 8-ribbed at least 1/2
length of capsule.
10. Capsule ovate to ovate-oblong, fully
exserted; exostome teeth 16.
11. Exostome fenestrate and cancellate; endemic
to
11. Exostome not fenestrate or cancellate, at
most perforate at tips; Arctic south to
10. Capsule
oblong to cylindric, emergent to slightly exserted;
exostome teeth 8.
12. Capsule oblong, ± 1/2 emergent; exostome
teeth perforate only at tips, not cancellate; endostome segments usually absent
when capsules are old and dry; leaves gradually narrowed to a long
acuminate-cuspidate, ± colorless apex; rare, western states, saxicolous . . 29.
Orthotrichum praemorsum
12. Capsule long-cylindric to oblong, 1/2
emergent to exserted; exostome teeth deeply perforate and cancellate, each
tooth appearing as split into 4 vertical rows; endostome segments usually
present when capsules are old and dry; leaves acute; common, throughout
northern North America, corticolous.
13. Capsule long-cylindric to cylindric, greater
than 1.5 mm, ribbed entire length, over 1/2 emergent; tapering gradually to
seta; endostome papillose; western
13. Capsule oblong to oblong-cylindric, less than
1.5 mm, ribbed for ± 1/2 length of capsule, ± emergent; tapering to seta
through a long, shriveled neck; endostome smooth; Great Lakes area,
northeastern North America, and Alaska . . 39 Orthotrichum sordidum
9. Exostome teeth recurved; endostome segments 8
or 16, large, stout, and thick, consisting of 2 rows of cells; capsule smooth
or weakly 8-ribbed no more than 1/2 length of capsule or if strongly 8-ribbed,
then dioicous.
14. Calyptra naked; endostome segments hyaline,
delicate, as wide as exostome teeth; capsule widest at mouth, ovate; leaves
lanceolate-oblong to oblong, rounded-obtuse; stems less than 1 cm, largest at
apex . . 12. Orthotrichum exiguum
14. Calyptra hairy; endostome segments yellowish,
stout, narrower than exostome teeth; capsule widest at middle or cylindric;
leaves lanceolate, acute or acuminate; stems longer than 1 cm, of uniform
thickness.
15. Dioicous; exostome teeth acuminate; capsule
oblong-cylindric, 8-ribbed . . 22. Orthotrichum lyellii
15. Autoicous; exostome teeth acute or truncate;
capsule of various types.
16. Leaves slenderly acute, tip contracted to
large, cuspidate point; peristome teeth truncate, often united at base to form
basal ring, endemic to Appalachian Mts. . . 18. Orthotrichum keeverae
16. Leaves acute or
acuminate; peristome teeth acute, separate to base; absent in Appalachian Mts.
17. Exostome teeth 16, endostome segments 16,
usually not erose;
17. Exostome teeth 8, or if 16 endostome erose;
endostome segments 8 to 16; temperate and boreal regions southward to
18. Capsule lightly 8-ribbed .
. 40. Orthotrichum speciosum
18. Capsule smooth.
19. Capsule cylindric,
exserted; exostome teeth 8; endostome segments 8 . .
10. Orthotrichum elegans
19. Capsule ovate, immersed; exostome teeth 16;
endostome segments 16 . . 45. Orthotrichum striatum
8. Endostome segments usually absent,
occasionally rudimentary; exostome teeth erect, sometimes reflexed when old and
dry; prostome usually present; leaves stiff, erect-appressed when dry, rarely
2-stratose.
20. Distal laminal cells 2-stratose, margins plane;
plants dark-green to black . . 6. Orthotrichum bolanderi
20. Distal laminal cells 1-stratose, margins
revolute; plants light- to olive-green.
21. Peristome rudimentary, appearing absent . . 41. Orthotrichum
spjutii
21. Peristome present, evident.
22. Exostome teeth smooth; leaves smooth;
calyptra naked; prostome large, extending 1/3 way up exostome; seasonally wet rocks . . 17. Orthotrichum
holzingeri
22. Exostome teeth papillose; leaves papillose;
calyptra hairy; prostome small or absent; dry rocks.
23. Capsule cylindric, fully exserted.
24. Exostome teeth usually 16, erect; capsule
smooth or slightly plicate; exothecial cells homogeneous; endostome
rudimentary, usually absent in open capsules . . 20. Orthotrichum laevigatum
24. Exostome teeth usually 8, reflexed-recurved;
capsule lightly 8-ribbed, exothecial cells differentiated for 1/2 length of
capsule; endostome usually present in at least some capsules
. . 34. Orthotrichum pylaisii (in part)
23. Capsule usually oblong and emergent.
25. Exostome teeth erect
to spreading; mostly on rocks, sometimes on trees . .
36. Orthotrichum rupestre
25. Exostome teeth recurved; on trees . . 37. Orthotrichum
shawii
7. Stomates immersed; basal leaf cells
rectangular, not nodose, thin-walled.
26. Leaves ± obtuse; margins coarsely dentate or
cucullate at apex; calyptra naked, distinctly plicate, papillose, peristome of
8, reflexed teeth; on seasonally wet rocks or bases of trees near streams,
sometimes submerged; west of continental divide.
27. Plants in tight, red-brown to black tufts;
leaf apices cucullate, entire to sometimes irregularly serrulate; costa
sinuose, often forked near apex; distal laminal cells to 20 \um wide . . 11. Orthotrichum euryphyllum
27. Plants in loose, dark-green to blackish-green
tufts; leaf apices flat, often dentate; costa straight; distal laminal cells to
13 \um wide . . 35.
Orthotrichum rivulare
26. Leaves ± acute or narrowly obtuse; margins
entire or denticulate near apex; calyptra naked or hairy, not distinctly
plicate, smooth; peristome various; on trunks of trees or dry rock faces;
variously distributed.
28. Capsules with 8 short ribs constricting
capsule at mouth when dry; exostome lacking; known only from
28. Capsules smooth or with 8 or 16 ribs
constricting capsule below mouth; exostome of 8 or 16 variably developed teeth;
throughout
29. Distal laminal cells uniformly
2-stratose.
30. Plants with all stems erect; leaves similar
on vegetative and fertile stems; scattered throughout western
30. Plants with creeping primary stems from which
erect vegetative and fertile stems arise; leaves of vegetative axes differentiated
from those of fertile axes; rare, restricted to
29. Distal laminal cells mostly 1-stratose,
occasionally 2-stratose in streaks, or with 2- to multistratose margins.
31. Exostome teeth erect or spreading, striate,
reticulate, densely papillose or papillose-striate; capsule with 8 or 16 ribs.
32. Endostome segments 16, linear, papillose, as
long as exostome. . 47. Orthotrichum
underwoodii
32. Endostome segments absent, or 8 and short,
not well-developed.
33. Capsule exserted, cylindrical, usually with 8
long and 8 short ribs; distal laminal cells 1-stratose .
. 4. Orthotrichum anomalum
33. Capsule immersed or emergent, rarely shortly
exserted, ovate or oblong, usually with 8 or 16 ± uniform ribs; distal laminal
cells 1- or partially 2-stratose.
34. Capsule with 16 ribs, ovate and constricted
below mouth when dry; leaves ovate-lanceolate, acute; distal laminal cells
1-stratose, rarely 2-stratose in streaks . . 8. Orthotrichum cupulatum
34. Capsule with 8 ribs, oblong or oblong-ovate,
sometimes constricted below mouth when dry; leaves lanceolate to ligulate,
obtuse or blunt; distal laminal cells 1- or 2-stratose.
35. Distal laminal cells of either laminae or
margins 2--7-stratose
36. Cells of leaf margins and of distal leaf
lamina mostly to completely 2-stratose . . 16. Orthotrichum hallii (in part)
36. Cells of leaf margins 2--7 stratose, laminal
cells 1-stratose, occasionally with a few 2-stratose streaks
. . 44. Orthotrichum strangulatum
35. Distal laminal cells 1-stratose.
37. Leaves appearing thick and glaucous; papillae
1--2, large, 2--3, forked; western
37. Leaves blackish green, dull; papillae 1--3, small,
conic; eastern and interior
38. Leaves oblong-lanceolate, margins plane or
nearly so; capsules oblong; exostome teeth reticulate-papillose. . 27. Orthotrichum parvulum
38. Leaves narrowly lanceolate, margins revolute;
capsules ovate; exostome teeth finely papillose. . 21. Orthotrichum lescurii
31.
Exostome teeth reflexed or recurved, finely papillose, rarely striate at
tips; capsule smooth or 8-ribbed.
39.
Capsule smooth (wrinkled and lightly 8-plicate when old) when dry; exostome
teeth 16, erect to recurved; endostome absent . . 32. Orthotrichum pusillum
39. Capsule 8-ribbed when dry; exostome teeth 8,
or 8 splitting to 16, reflexed; endostome usually present
40. Leaf apex cuspidate, usually denticulate, margins
incurved and forming channel at apex; southwestern U.S.A . . .5.
Orthotrichum bartramii
40. Leaf apex obtuse, acute, or apiculate,
margins not forming channel at apex; throughout
41. Stomates scarcely to
1/2 covered by non-projecting subsidiary cells, inner walls not much thickened;
calyptra usually naked, sometimes with a few hairs, oblong to conic-oblong.
42. Leaves acute,
narrowly obtuse, or apiculate-crenulate, oblong-lanceolate to lanceolate;
distal laminal cells less than 14 \um; calyptra conic-oblong
. . 26. Orthotrichum pallens
42. Leaves apiculate,
mucronate, or rounded-obtuse, ovate or narrowly ovate-lanceolate; distal
laminal cells greater than 13 \um; calyptra oblong.
43. Leaves ovate to ovate-oblong, rounded-obtuse,
at least some leaves mucronate, carinate; western
43. Leaves ovate-lanceolate to lanceolate, acute,
most leaves apiculate, not carinate; throughout
41. Stomates 1/2 to completely covered
by projecting subsidiary cells, inner walls thickened; calyptra usually with a
few to many hairs, oblong.
44. Leaves with 2- to 3-stratose margins; dry
granite rocks . . 38. Orthotrichum shevockii
44. Leaves with 1-stratose margins; trees and
mesic rocks.
45. Exostome teeth
striate-reticulate in distal portion; calyptra with strongly papillose hairs;
leaf cells with regular, long-forked papillae; leaves slightly twisted when
dry; capsule strangulate when dry. . . 2.
Orthotrichum alpestre
45. Exostome teeth papillose; calyptra hairs
smooth or papillae formed by projecting cell walls; leaf cells smooth or with
low, conical papillae, never forked; leaves stiff, erect-appressed when dry; capsule
variously constricted beneath mouth when dry.
46. Vaginula with numerous hairs.
47. Capsules with thin, delicate ribs when dry;
47. Capsules with thick, broad ribs when dry;
46. Vaginula without hairs.
48. Leaf tips incurved when dry; capsule
cylindric to cylindric-elliptic,
contracted along entire length, not constricted beneath mouth; stomates
in proximal 1/3 of capsule, completely covered by subsidiary cells; restricted
to west coast . . 46. Orthotrichum tenellum
48. Leaf tips not incurved when dry; capsule
oblong-cylindric, sometimes constricted beneath mouth, never contracted along entire
length; stomates at middle of capsule, completely covered by subsidiary cells
or not;
49. Leaf cells greater than 14 \um, lumina smooth
and rounded; leaves apiculate or slenderly acute, ovate-lanceolate to
oblong-lanceolate; subsidiary cells not completely covering stomates
. . 31. Orthotrichum pumilum
49. Leaf cells less than 14 \um, lumina irregular
due to uneven wall thickenings; leaves broadly acute to obtuse, lanceolate;
subsidiary cells completely covering stomates.
50. Capsule not
constricted beneath mouth, yellow, with thin, delicate ribs; exostome teeth 16,
rarely united into 8 pairs; subsidiary cells projecting upward over stomate . . 25. Orthotrichum
ohioense
50. Capsule strongly constricted beneath mouth,
brown, with thick, broad ribs; exostome teeth 8; subsidiary cells projecting
horizontally over stomate . . 42. Orthotrichum stellatum
