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BFNA Title: Orthotrichum |
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ORTHOTRICHUM -
ORTHOTRICHACEAE 6. Orthotrichum
Hedwig, Spec. Musci. 162. 1801 * [Named for the straight, erect hairs
found on the calyptra in many of the 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 leaf cells mostly rounded, almost
isodiametric, usually 6--24(--30) µm, 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 120 (40
in the flora); worldwide, more common in temperate areas. SELECTED
REFERENCES: Lewinsky, J. 1993.
Monographic studies on Orthotrichum
(Musci). Bryobrothera, Vol. 2:
1--59. Vitt, D. H. 1973.
A Revision of the Genus Orthotrichum
in North America, North of Mexico.
Bryophytorum Bibliotheca, 1: 1--108. 1.
Leaves ending in ± serrate, hyaline awn . . 8. Orthotrichum diaphanum 1. Leaves obtuse, acute, acuminate, or
cuspidate, not ending in hyaline awn. 2.
Leaf margins erect, incurved, or involute; dioicous; leaves ovate or
elliptic, with broadly obtuse or rounded apices; calyptrae papillose, not
plicate or hairy. 3.
Leaf margins involute; distal leaf cells with 2--3 conical papillae
per cell; peristome lacking; Newfoundland . . 13. Orthotrichum gymnostomum 3.
Leaf margins erect-incurved to incurved; distal leaf cells with 1
papilla per cell; peristome double; common throughout boreal and western
regions . . 19. Orthotrichum
obtusifolium 2.
Leaf margins plane, recurved, revolute, or thickened; condition autoicous or if dioicous then
leaves acuminate; leaves ± lanceolate or oblong, with narrowly obtuse, acute,
or acuminate apices; calyptra smooth or rarely papillose, plicate, hairy, or
naked. 4.
Leaves crisped-flexuose when dry; capsule cylindric or
elliptic-cylindric, fully exserted; stomates immersed; Pacific Northwest and
Alaska. 5.
Plants 0.8--2 mm; distal leaf cells 7--11 µm wide; seta usually 2.5--6
mm; capsule long-exserted, ovate-cylindric when mature; peristome teeth 8,
light-brown, finely papillose on the exterior, finely striate on the
interior; stomates immersed, in the middle and proximal portion of the
capsule . . 6. Orthotrichum consimile 5.
Plants usually less than 1.3 cm; distal leaf cells 9--14 µm wide; seta
1.5--3.5 mm; capsule barely exserted, oblong-cylindric when mature; peristome
teeth 16, red, irregularly papillose-reticulate; stomates immersed above the
middle of the capsule, never in the proximal portion . . 24. Orthotrichum pulchellum 4.
Leaves erect-appressed when dry; capsules various; stomates
superficial or immersed; throughout North America. 6.
Stomates superficial; basal leaf cells elongate, ± nodose,
thick-walled. 7.
Endostome segments well-developed, usually present when capsule is
old; exostome teeth recurved or reflexed; prostome never present; leaves
sometimes flexuose when dry, always 1-stratose 8.
Exostome teeth reflexed; endostome segments 8, narrow, consisting of a
single row of cells; capsule strongly 8-ribbed at least 1/2 length of
capsule. 9.
Exostome fenestrate and cancellate; endemic to St. Paul's Island,
Alaska . . 11. Orthotrichum fenestratum 9.
Exostome not fenestrate or cancellate, at most perforate at tips;
Arctic south to California . . 28. Orthotrichum
pylaisii 8.
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. 10.
Calyptra oblong-conic, naked; endostome segments hyaline, delicate, as
wide as exostome teeth; capsule widest at mouth, ovate; leaves oblong-ovate,
rounded-obtuse; plants never longer than 1 cm, largest at apex . . 10. Orthotrichum exiguum 10.
Calyptra short-conic, hairy; endostome segments yellowish, stout, not
as wide as exostome teeth; capsule widest at middle or cylindric; leaves
lanceolate, acute or acuminate; stems longer than 1 cm, of uniform thickness. 11.
Plants dioicous; exostome teeth acuminate; capsule long-cylindric and
8-ribbed . . 18. Orthotrichum lyellii 11.
Plants autoicous; exostome teeth acute or truncate; capsule of various
types. 12.
Leaves slenderly acute, tip contracted to large, cuspidate point; peristome
teeth truncate, often united at base to form basal ring, endemic to
Appalachian Mts. . . . 16. Orthotrichum
keeverae 12.
Leaves acute or acuminate; peristome teeth acute, separate to base;
not found in Appalachian Mts. 13.
Exostome teeth 16, endostome segments 16, usually not erose; New
Mexico and Arizona . . 27. Orthotrichum
pycnophyllum 13.
Exostome teeth 8, or if 16 endostome erose; endostome segments 8 to
16; temperate and boreal regions southward to California and Missouri. 14.
Capsule lightly 8-ribbed . . 33. Orthotrichum
speciosum 14.
Capsule smooth. 15. Capsule cylindric, exserted;
exostome teeth 8; endostome segments 8 . . 9. Orthotrichum elegans 15.
Capsule ovate, immersed; exostome teeth 16; endostome segments 16. . .
38. Orthotrichum striatum 7.
Endostome segments rudimentary or, more often, absent; exostome teeth
erect, sometimes reflexed when old and dry; prostome usually present; leaves
stiff, erect-appressed when dry, rarely 2-stratose 16.
Leaves 2-stratose, margins plane; plants dark-green to black . . 5. Orthotrichum bolanderi 16.
Leaves 1-stratose, margins revolute; plants light- to olive-green. 17.
Peristome rudimentary, appearing absent . . 34. Orthotrichum spjutii 17.
Peristome present. 18.
Exostome teeth smooth; leaves smooth; calyptra naked; prostome large,
extending 1/3 way up exostome; seasonally wet rocks . . 15. Orthotrichum holzingeri 18.
Exostome teeth papillose; leaves papillose; calyptra hairy; prostome
small or absent; dry rocks. 19.
Capsule cylindric, fully exserted. 20.
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 .
. 28. Orthotrichum pylaisii 20.
Exostome teeth usually 16, erect; capsule smooth or slightly plicate;
exothecial cells not or scarcely differentiated; endostome rudimentary,
usually absent in open capsules . . 17. Orthotrichum
laevigatum 19. Capsule usually oblong and emergent, if
cylindric and exserted then strongly 8-ribbed. 21.
Exostome teeth erect; endostome absent . . 30. Orthotrichum rupestre 21.
Exostome teeth reflexed; endostome present. 22.
Capsule ovate to ovate-oblong, fully exserted; exostome teeth 16. 23.
Exostome fenestrate and cancellate; endemic to St. Paul's Island,
Alaska . . 11. Orthotrichum fenestratum 23.
Exostome not fenestrate or cancellate, at most perforate at tips;
Arctic south to California . . 28. Orthotrichum
pylaisii 22. Capsule oblong
to cylindric, emergent to slightly exserted; exostome teeth 8. 24.
Capsule oblong, ± 1/2 emergent; exostome teeth perforate only at tips,
not cancellate; endostome segments usually not present when capsules are old
and dry; leaves gradually narrowed to a long acuminate-cuspidate apex, which
is ± colorless; rare, western states, saxicolous . . 23. Orthotrichum praemorsum 24.
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. 25.
Capsule long-cylindric to cylindric, greater than 1.5 mm, ribbed entire
length, emergent over 1/2; tapering gradually to seta, which is usually over
1 mm in length; endostome papillose; western North America . . 1. Orthotrichum affine 25.
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, which is less than 1.5 mm in length; endostome smooth; Great Lakes
area, northeastern North America, and Alaska . . 32. Orthotrichum sordidum 6.
Stomates immersed; basal leaf cells rectangular, not nodose,
thin-walled. 26.
Leaves ± obtuse; margins coarsely dentate near apex; calyptra naked,
distinctly plicate, papillose, peristome of 8, reflexed teeth; rocks or bases
of trees near streams, sometimes submerged; west of continental divide. . 29.
Orthotrichum rivulare 26.
Leaves ± acute or narrowly obtuse; margins entire or denticulate near
apex; calyptra naked or hairy, not distinctly plicate, smooth; peristome
various; trunks of trees or dry rock faces; variously distributed. 27.
Exostome teeth erect or spreading, striate, reticulate, or
papillose-striate; capsule with 8 or 16 ribs. 28.
Endostome segments 16, linear, papillose, as long as exostome. . 40. Orthotrichum underwoodii 28.
Endostome segments absent, or 8 short, not well-developed. 29.
Capsule exserted, cylindrical, usually with 8 long and 8 short ribs;
leaves 1-stratose . . 3. Orthotrichum
anomalum 29.
Capsule immersed or emergent, rarely shortly exserted, ovate or
oblong, usually with 8 or 16 ± uniform ribs; leaves 1- or 2-stratose. 30.
Capsule with 16 ribs, immersed to 1/3 emergent, ovate and constricted
below mouth when dry; leaves ovate-lanceolate, acute, 1-stratose or rarely with
2-stratose streaks . . 7. Orthotrichum
cupulatum 30.
Capsule with 8 ribs, ± emergent, oblong or oblong-ovate, sometimes
constricted below mouth when dry; leaves lanceolate to ligulate, obtuse or
blunt, 1- or 2-stratose. 31.
Exostome teeth finely papillose or finely reticulate-papillose, 16,
yellowish, erect or spreading-recurved; leaf margins 1--5 cells thick, at
least in some leaves thicker than lamina; capsule usually strongly
constricted below mouth when old; eastern North America . . 37. Orthotrichum
strangulatum 31.
Exostome teeth ridged-striate, coarsely striate-reticulate, or
coarsely papillose-striate, 8 splitting to 16, white to clear, incurved,
spreading, or reflexed; leaf margins not thicker than lamina; capsule only moderately
constricted below mouth when dry; western North America. 32.
Leaves 2-stratose or partially 2-stratose, green, lanceolate, obtuse,
loosely-appressed, not incurved when dry; papillae small and conical;
exostome teeth usually 8 . . 14. Orthotrichum
hallii 32.
Leaves 1-stratose, usually thick and glaucous, ligulate, blunt, stiff,
incurved when dry; papillae large, 2-3 forked; exostome teeth ± 16. . . 22. Orthotrichum
pellucidum 27.
... Exostome teeth
reflexed or recurved, papillose, rarely striate at tips; capsule smooth or
8-ribbed. 33.
.... Capsule smooth
(wrinkled and lightly 8-plicate when old) when dry; exostome of 16, erect to
recurved teeth; endostome absent . . . Orthotrichum
pusillum 33.
Capsule 8-ribbed when dry; exostome teeth 8, or 8 splitting to 16, reflexed;
endostome usually present 34.
Leaf apex cuspidate and usually denticulate, margins incurved and
forming channel at apex; southwestern U.S.A. . .4. Orthotrichum bartramii 34.
Leaf apex obtuse, acute, or apiculate, margins not forming channel at
apex; throughout North America. 35. Stomates scarcely to 1/2 covered by
non-projecting subsidiary cells, inner walls not much thickened; calyptra usually naked or
sometimes with a few hairs, oblong to conic-oblong. 36. Leaves acute, narrowly obtuse, or
apiculate-crenulate, oblong-lanceolate to lanceolate; distal leaf cells less
than 14 µm; calyptra conic-oblong . . 21. Orthotrichum
pallens 36. Leaves apiculate, mucronate, or
rounded-obtuse, ovate or narrowly ovate-lanceolate; distal leaf cells greater
than 13 µm; calyptra oblong. 37.
Leaves ovate to ovate-oblong, rounded-obtuse, at least some leaves
mucronate, carinate; western North America . . 12. Orthotrichum flowersii 37.
Leaves ovate-lanceolate to lanceolate, acute, most leaves apiculate,
not carinate; throughout North America . . 25. Orthotrichum pumilum 35.
Stomates 1/2 to completely covered by projecting subsidiary cells,
inner walls thickened; calyptra usually with a few to many hairs, oblong. 38.
Leaves with 2- to 3-stratose margins; dry granite rocks . . 31. Orthotrichum shevockii 38.
Leaves with 1-stratose margins; trees and mesic rocks. 39. Exostome teeth with
striate-reticulate ornamentation in distal portion; calyptra with strongly
papillose hairs; leaf cells papillose, with regular, long-forked papillae;
leaves slightly twisted when dry; capsule strangulate when dry. . . 2. Orthotrichum
alpestre 39.
Exostome teeth papillose; calyptra hairs smooth or papillae formed by
projecting cell walls; leaf cells smooth or with low, conical papillae,
papillae never forked; leaves stiff, erect-appressed when dry; capsule
variously constricted beneath mouth when dry. 40.
Vaginula with numerous hairs . . 36. Orthotrichum stramineum 40.
Vaginula without hairs. 41.
Leaf tips incurved when dry; capsule cylindric to cylindric-elliptic,
not constricted beneath mouth, but contracted along entire length; stomates
in proximal 1/3 of capsule, completely covered by subsidiary cells;
restricted to west coast . . 39. Orthotrichum tenellum 41.
Leaf tips not incurved when dry; capsule oblong-cylindric, constricted
beneath mouth, never contracted along entire length; stomates at middle of
capsule; Rocky Mountains and eastward. 42.
Leaf cells greater than 14 µm, the lumina smooth and rounded; leaves
apiculate or slenderly acute, ovate-lanceolate to oblong-lanceolate;
subsidiary cells not completely covering stomates . . 39. Orthotrichum pumilum 42.
Leaf cells less than 14 µm, lumina irregular due to uneven wall
thickenings; leaves broadly acute to obtuse, lanceolate; subsidiary cells
completely covering stomates. 43.
Capsule strongly constricted beneath mouth, brown; exothecial cells
with thickened walls, in rows 6--8 cells wide; exostome teeth 8; subsidiary
cells projecting horizontally over stomate . . 35. Orthotrichum stellatum 43. Capsule not constricted beneath
mouth, yellow; exothecial cells with thickened walls in rows 4--6 cells wide;
peristome teeth 16, rarely united into 8 pairs; subsidiary cells projecting
upward over stomate . . 20. Orthotrichum
ohioense 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 µm 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 µm. 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 the western
portion of North America, is best recognized by the following combination of
characters: superficial stomates;
cylindric, strongly ribbed, almost exserted capsules; a well-developed, but
not stout endostome; and a calyptra that is only sparsely hairy. 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 µm 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 µm. 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 leaf cells, slightly twisted
leaves, which are always sharply pointed, a calyptra with papillose hairs,
the absence of a preperistome, and the papillose or papillose-reticulate
exostome. The endostome is always
present in mature capsule and because of the enlarged crosswalls has a shiny
appearance. The capsule, as it is
usually seen, is oblong and strangulate, although the shape depends on age,
and in some specimens the capsule is short-cylindric and not much constricted
under the mouth. 3. Orthotrichum
anomalum Hedwig, Spec. Musc. 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 µm 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 µm. 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
1-stratose leaves. The most useful
character in distinguishing this species from others is the exserted capsule
with 8 long ribs alternating with 8 shorter ones. 4. 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
µm 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 µm. Bark of Quercus or Populus tremuloides; ±
3000 m; Ariz.; Mexico; Central America (Guatemala). The channeled,
cuspidate apex of the mature leaves distinguishes this species 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 5. Orthotrichum
bolanderi Sullivant, Icon. Musc.
Suppl. 64 Pl. 46. 1874 Plants 0.5--4 cm. 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 µm 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 µm. Rocks,
especially sandstone; 100--2000 m;
Calif., Wash.; Mexico; Central America (Guatemala). Orthotrichum bolanderi
is the only species with superficial stomates that has 2-stratose leaves. It
has leaves with a subsheathing base, ovoid cylindric capsules with a long
wrinkled neck, and exostome teeth that are reflexed-recurved. When dry the leaves are stiffly
erect-appressed and when wet they lie flat on a microscope slide. 6. 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 contorted-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 µm 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 mature, 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 µm. Tree trunks
and branches in coniferous forests, sometimes found on boulders in shaded
habitats; low elevations; B.C.; Alaska, Calif., Idaho, Oreg., Wash.; Europe. 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. 7. 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 µm 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 µm. 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 Africa. Orthotrichum cupulatum
is extremely variable but can be distinguished by the following combination of
characters most of which are usually evident:
(1) leaves broadly but sharply
acute, ovate-lanceolate, and usually 1-stratose or occasionally partially
2-stratose; (2) papillae of the leaf cells usually large
and forked, but if small and conical then the leaf is 1-stratose; (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. 8. 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 µm 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 µm. 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 that lack 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. 9. 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 µm 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 µm. 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 forming 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. 10. 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 µm 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 µm. Base of trees
or on tree trunks; 10--200 m; Ga., N.C., S.C., Tenn., Va.; e Asia. The distal leaf
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. 11. 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 µm wide, 1-stratose, with 1--2 small conical papillae per
cell; basal laminal cells narrowly rectangular, grading to subquadrate at
margin, nodose. 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, 1 row of cells, papillose-reticulate; exostome teeth 8
splitting to 16, erect to recurved when old, coarsely papillose above,
becoming finely papillose to almost smooth near base. Calyptra
short-conic, plicate, with few, thick, white hairs. Spores
20--27 µm. Collected only
once, by J. M. Macoun, who did not record the substrate; probably near sea
level; Alaska. Endemic to Alaska. The exostome
teeth that are cancellate to the base and fenestrate and perforate only in
the distal portion, and the ovate capsule, distinguish this species. 12. 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 µm 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 µm. Base of tree
trunks and trunks of deciduous trees in xeric and exposed habitats; 500--2000 m; Calif., Idaho, Nev., Utah;
Europe. This species
is distinguished from O. pumilum and O. pallens by having
blunt, carinate leaves that are ovate or ovate-oblong, and almost always have
a small mucro. 13. 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 µm 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 µm. Bark of
deciduous trees; 0--200 m; Nfld.;
Europe; Asia (Japan). This species
differs from O. obtusifolium (a species with similar
appearance) by having leaves with involute margins, cells with 2--3 papillae,
and immersed capsules with no peristome, while O. obtusifolium has
leaves with erect margins, unipapillose cells, and emergent capsules with a double peristome. Orthotrichum
gymnostomum is rare in North
America, restricted to the coastal areas of Newfoundland. 14. Orthotrichum
hallii Sullivant & Lesquereux
in W. S. Sullivant, Icones Musc. Suppl. 63.
Pl. 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 µm wide, 2-stratose, rarely with a few 1-stratose 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 µm. 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 North America. Orthotrichum hallii
is distinguished by having leaves that are completely, or nearly so,
2-stratose in their distal portion, and oblong, 8-ribbed capsules that are
emergent. 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 in all
proportions. The papillae are always
small and conical. 15. 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 µm 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 µm. Seasonally wet
rocks in small streams of dry montane forests; 1000--2000 m; Calif., Idaho,
Oreg., Wash., Wyo. Restricted to w
North America This species
is distinguished by having smooth leaf cells, smooth exostome, naked
calyptra, and exserted, lightly-ribbed capsules. It is a distinct species, endemic to a
small region in western North America, and is most closely related to O. laevigatum. 16. 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; margins revolute nearly to apex, often irregularly notched
near apex; distal laminal cells 7--14 µm 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. Calyptra
conic-oblong, with papillose hairs 1--3 cells thick. Spores
17--27 µm. Trunks and
branches of live and dead Quercus
montana; ± 1500--2000 m; N.C., Va.
Endemic. Orthotrichum keeverae
is the only species in southeastern North America with 16, recurved, exostome
teeth; a stout endostome and cuspidate leaf apices. The exostome teeth are truncate and
partially fused at the base forming a basal membrane. 17. 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 µm 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 16, erect, papillose-granulose, coarsely
papillose or striate-papillose. Calyptra conic-oblong, sparsely
hairy, hairs finely papillose. Spores 10--15 µm. Usually on
non-calcareous boulders in mesic, yet 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. 18. Orthotrichum
lyellii Hooker & Taylor, Musc.
Brit. 76. 22. 1818 Dorcadion lyellii
(Hooker & Taylor) Lindberg; Orthotrichum
lyellii var. howei Renault & Cardot; O.
lyellii var. papillosum (Hampe)
Lesquereux & James; O. lyellii
var. pringlei (J. K. A. Müller)
Macoun & Kindberg; O. lyellii
ssp. strictum Venturi; O. lyellioides Kindberg; O. menziesii Mitten; O. papillosum Hampe; O. papillosum ssp. strictum (Venturi) Kindberg; O. pringlei J. K. A. Müller 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 µm 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, distinctly 8-ribbed from 1/2 to entire length; stomates superficial; peristome double; endostome segments 16, large, coarsely papillose; exostome teeth 16, reflexed-recurved, densely papillose. |