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VOL. XII, No. 3, PP 271-328. FIGS. 1-30. AUGUST 15, 1910
THE POLYTRICHACE.E OF WESTERN NORTH
AMERICA
BY T. C. FRYE
University of Washington, Seattle, Washington
INTRODUCTORY NOTE.
There is great confusion in the taxonomy of our western mosses on
account of the duplication of names and the naming of species from
sterile specimens or single collections. Systematic work on them is
much needed. To show the way, a single family, the Polytricha-
ceae, was studied from type material and accessible collections. In
the keys in this paper the characters separating the genera or
species are given for all, thus making a comparison. So often one
wants a comparison rather than a description, that it is hoped this
will prove of value to those using this paper.
Acknowledgments are due to Mrs. Elizabeth G. Britton for kindly
permitting the writer to examine type material in the herbarium of
the New York Botanical Gardens.
The illustrations in this paper are mostly from drawings by Elsie
K. Waddingham.
POLYTRICHACE^E.
Name derived from poly = many, and tricho = hair; referring to
the hairiness of the calyptra in many genera.
Plants usually of large size, growing on soil. Stems simple or
slightly branched, springing from a subterranean shoot.
Leaves usually narrow. Lamellae present on their inner surface
and sometimes on the back as well, each usually a few cells high and
Proc. Wash. Acad. Sci., August, 1910.
272 FRYE
i cell thick, wanting in Racelopus; marginal cells often of a different
form from the others. Vein one, extending at least nearly to the tip.
Inflorescence nearly always dioicous; sex organs terminal; antheridia
in a rather large cup or disk through which the plant generally again
grows.
Calyptra narrow, often covered with a dense mat of branched
hairs which are directed away from the point and end freely; if not
with densely matted hairs, the apex of the calyptra is spinulose, or
has few to many hairs, or is quite smooth.
Capsule large, cylindric, or prismatic with 2 — 6 angles. Peri-
stome present (except in Lyellia, Bartramiopsis and some species of
Psilopilum), single; teeth 32 or 64, rarely 16, unbarred, ligulate, tri-
angular in cross section. Columella expanded at the apex into a
shield-shaped membrane (epiphragm) covering the mouth of the cap-
sule and uniting with the tips of the teeth. Lid present. Pedicel
long, smooth (except in Racelopus). Spores .008 — .021 mm., rarely
larger, smooth or nearly so.
Total number of genera, 10; number.represented in western North
America, 7. Total number species, about 320; number in western
North America, 27.
SYNOPSIS AND COMPARISON OF THE WORLD'S GENERA OF POLYTRI-
CHACE^E.
i. Calyptra with few or no hairs.
2. Lamellae on upper side 18 or fewer.
3. Lamina of i layer of cells.
4. Leaves bordered.
5. Peristome present.
6. Stomates wanting on capsules.
7. Capsule terete Catharinea.
4. Leaves not bordered.
5. Peristome present; calyptra with few hairs; lamellae on
vein at back of leaves in nearly all cases.
6. Stomates present on capsule.
7. Capsule terete Oligotrichum.
5. Peristome wanting; or if present, calyptra without hairs ;
without lamellae on back of leaves.
6. Stomates present on capsule.
7. Capsule somewhat flattened, elliptical in cross section.
Psilopilum.
3. Lamina composed of 2 layers of cells.
4. Leaves not bordered.
THE POLYTRICHACE^E OF WESTERN NORTH AMERICA 273
5. Peris tome wanting.
6. Stomates present on capsule.
7. Capsule terete Bartramiopsis.
5. Peris tome present.
6. Stomates present on capsule.
7. Capsule terete Dendroligotrichum.
2. Lamallae on upper side 20 or more.
3. Lamina of i layer of cells.
4. Leaves not bordered.
5. Peristome wanting.
6. Stomates present on capsule.
7. Capsule 2-4 angled Polytrichadelphus.
3. Lamina of 2 layers of cells. .
4. Leaves not bordered.
5. Peristome wanting.
6. Stomates present on capsule.
7. Capsule keeled on one side, somewhat oval in cross
section Lyellia.
i. Calyptra densely covered with hairs.
2. Lamellae wanting.
3. Lamina of i layer of cells.
4. Leaves not bordered.
5. Peristome present.
6. Stomates wanting on capsule.
7. Capsule terete Racelopus,
2. Lamellae on upper side 20 or more.
3. Lamina of i layer of cells.
4. Leaves not bordered.
5. Peristome present.
6. Stomates either present or wanting on the capsule.
7. Capsule terete Pogonatum.
6. Stomates present on capsule.
7. Capsule 4-6 angled1 Polytrichum.
'The writer has followed the classification of Brotherus in Engler & Prantl: "Die
natiirlichen Pflanzenfamilien, " Teil i, Abt. 3, s. 669-698 (1904) except in the separa-
tion of Pogonatum from Polytrichum. The chief question here arising is where to
put Pogonatum alpinum. Excepting this plant the Pogonatums have capsules terete,
teeth 32, stomates wanting; the Polytrichums have capsules angular, teeth 64, stomates
present; Pogonatum alpinum is a fine gradation form between the two genera, having
the terete capsule of the Pogonatums, the stomates of the Polytrichums, and standing
between the two genera in its 32 or 64 teeth depending upon how one counts the doub-
ling. Whether one classes it a Pogonatum or a Polytrichum depends upon whether one
emphasizes the form of the capsule or the presence of the stomates. When there is
nothing to be gained by using a microscopic character in the separation of genera,
why not use one which can be seen with the naked eye? In this paper Pogonatum
alpinum is therefore classed as a Pogonatum.
274 FRYE
KEY TO THE WEST NORTH AMERICAN GENERA.
i. Lamellae on upper side of leaf 18 or fewer 2.
1. Lamellae on upper side of leaf 20 or more 8.
2. Leaves bordered Catharinea, p. 275
2. Leaves not bordered 3.
3. Leaf margin with long hairs where sheath grades into blade;
lamina of two layers of cells; peristome wanting.
Bartramiopsis, p. 289
3 . Leaf margin without hairs ; lamina of i layer of cells ; peristome
present 4.
4. Capsule distinctly cernuous Psilopilum, p. 288
4. Capsule straight or very nearly so Oligotrichum, p. 281
4. Capsule wanting (for sterile specimens) 5.
5. Lamellae straight; plant much crisped when dry.
Oligotrichum parallelum, p. 282
5. Lamellae wavy from side to side ; plants not or very little crisped
when dry 6.
6. Lamellae on back conspicuous, about equal in number and size
to those on the upper side; leaf margin plane or erect.
Oligotrichum aligerum, p. 284
6. Lamellae on back inconspicuous or none ; leaf margin incurved. 7.
7. Leaf margin abruptly incurved above the sheath; lamellae on
back inconspicuous or none. .Oligotrichum incur vum, p. 285
7. Leaf margin gradually incurved from base; lamellae on back
wanting Psilopilum, p. 298
8. Capsules 4-6 angled; teeth 64; calyptra densely hairy.
Polytrichum, p. 304
8. Capsules 2-4 angled ; two of the angles nearer together than the
others; teeth 64, calyptra with few hairs.
Polytrichadelphus, p. 291
8. Capsules terete ; teeth 3 2 , or 64 in Pogonatum alpinum; calyptra
densely hairy Pogonatum, p. 294
8. Capsules wanting (for sterile specimens) 9.
9. Leaf margin entire, inflexed over the lamellae in most species.
Polytrichum, p. 304
9. Leaf margin serrate 10.
10. Leaves very much crisped when dry.
Pogonatum contortum, p. 295
10. Leaves little or not at all crisped when dry n.
11. Marginal cells of lamellae smooth or merely grooved.
Polytrichum, p. 304
ii. Marginal cells of lamellae rough or papillose 12.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 275
12. Plants erect; marginal cells of lamellae not higher than wide,
except in P. alpinum Pogonatum, p. 294
12. Plants decumbent; marginal cells higher than wide.
"Polytrichadelphus, p. 291
CATHARINEA Ehrh.
Atrichum Beauv.
Named after the Empress Catharine II of Russia.
Plants not very tall. Stems with abundant rhizoids at base.
Leaves without sheath, more or less transversely undulate, gener-
ally with diagonal rows of teeth on the back, mostly crisped when
dry. Margin plane, bordered except in young leaves, singly or
doubly serrate with coarse sharp teeth. Lamellae i-io, restricted
to the vein, composed of similar smooth cells, margin entire. Vein
f- the leaf-width or less, percurrent or vanishing, without lamellae on
the back, often toothed on back near apex. Cells all smooth, chloro-
phyllose, mostly quadratic, upper ones hexagonal or elliptic-hexa-
gonal.
Calyptra naked, or with a few hairs or teeth at the tip.
Capsule somewhat inclined, curved, never angular, smooth, with-
out stomates. Peristome present; teeth 32, pale, with yellow or
brown middle line. Lid long-beaked. Pedicel smooth, solitary or
sometimes 1-3 from a stem-tip.
Number species in western North America, 4; total number
species, about 33.
THE WEST NORTH AMERICAN SPECIES, — A COMPARISON AND KEY.
i . Capsule as 1 14 ; leaves not wavy along the margin when moist.
2. Vein constituting J-^ of the leaf -width.
3. Lamellae 1-2 cells high.
4. Cells .025-. 045. mm.
5. Lamina without teeth at back /. C. crispa.
i . Capsule as i : 6-8 ; leaves slightly to distinctly wavy along mar-
gins when moist.
2. Vein constituting J-J of the leaf -width.
3. Lamellae 5-8 cells high.
4. Cells . OIO-.O2O mm.
5. Lamina mostly with teeth at back 2. C. angustata.
2. Vein constituting J-j | of leaf- width.
3. Lamellae 3-5 cells high.
276 FRYE
4. Cells .OIO-.O2O mm.
5. Lamina mostly with teeth at back 3. C. undulata.
3. Lamellae 9-13 cells high.
4. Cells .025-. 050 mm.
5. Lamina mostly with teeth at back 4. C. selwyni.
1. Catharinea crispa James, in Proc. of Amer. Acad., 1855, p. 445.
Atrichum crispum Sull., in Mos. of U. S., p. 41 (1856).
Name probably derived from the crisping of the leaves in drying,
a characteristic however, not restricted to this species, nor even this
genus.
Plants dioicous, 2.5-10 cm. high, in soft tufts. Stems erect, sim-
ple.
Leaves distant, oval-oblong to oblong-lanceolate, not papillose,
patent, crisped when dry, obtusely acuminate, hardly at allundulate,
width to length about as i: 2-3, the lower shorter and broader;
lamina smooth at back. Margin reddish, toothed from near the
base, bordered. Lamellae 1-4, 1-3 cells high, vanishing in the lower
half of the leaf. Vein strong, reddish/brown, vanishing in the apex,
with few or no teeth at back, about i-yV the leaf -width. Cells .025-
.045 mm., quadrate-hexagonal or rounded, the lower elongate.
Calyptra smooth except at tip, which is roughened with very short
hairs.
Capsule erect or nearly so, slightly curved, width to length about
as 1:4, narrowly obconic, wide-mouthed. Teeth narrow, unequal;
basal membrane very narrow. Lid conic, with subulate beak.
Pedicels slender, 1-3 at a stem-tip, somewhat flexuose. — On clayey
soil. — Revelstoke, British Columbia; Atlantic Coast of United
States; England.
2. Catharinea angustata Brid., in Mant. Muse. p. 204 (1819),
Atrichum angustatum B. & S., in Bryol. Eur. t. 411, (1844.)
Name derived from augustus = narrow; referring to the narrow
leaves.
Plants dioicous. Stems 2-5 cm. high.
Leaves narrower than in C. undulata, undulate when moist, width
to length about as i: 7-10, not papillose; lamina smooth at back.
Margin serrate only above middle, bordered. Damellae 4-7, 5-8
cells high. Vein toothed at back, \-\ the width of leaf. Cells
.010 — .014 mm.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
277
Calyptra cucullate, about half covering the capsule, rough at tip.
Capsule purple, narrower and more erect than in C. undulata,
width to length about as i: 7-8. Teeth shorter than in C. undu-
lata. Lid dark purple, about half as long as the capsule. — On
clayey soil. — McLeod's Lake, British Columbia; Santa Cruz
Mountains, California; Atlantic states; Eastern Canada; Europe.
Fig. 1. Catharinea crispa.
1 = Plant moist, with capsule, X i. 2 = Plant dry, showing crisping of leaves
X i. 3 = Capsule with lid, X 5. 4 = Peristome X 150. 5 = Leaf showing lamel-
lae on upper side, X 15. 6 = Leaf- tip, X 65. 7 = Cross section of leaf, showing
lamellae, X 65. 8 = Cross section of leaf margin showing thickened border cells,
X 250. 9 = Cross section of a few lamellae, X 250.
3. Catharinea undulata Web. & Mohr, in Ind. pi. crypt. (1803).
Atrichum undulatum Beauv. Prodr. p. 42 (1805).
So named on account of its wavy or undulate leaves.
Plants in loose patches, dull green. Stems erect, 2.5-5 cm-
high, simple or much branched, from a subterranean shoot. Inflor-
escence autoicous.
278
FRYE
Lower leaves very small, scale-like; upper leaves ligulate, width to
length about as i : 5-6, 4-6 cm. long, strongly transversely undu-
late, much crisped and incurved when dry, spreading when moist,
lamina with transverse rows of teeth on back. Margin bordered
with 2 — 3 rows of very narrow brownish cells, sharply spinose for the
greater part of its length with strong and usually paired teeth.
Lamellae 2-6, straight, 3-5 cells high, occasionally as much as 7
Fig. 2. Catharinea angustata.
1 = Plant dry, X i. 2 = Moist plant, with capsule, X i. 3 = Capsule, X 5.
4 = Peristome, X 150. 5 = Leaf showing lamellae on upper side, X 15. 6 = Leaf
tip, X 65. 7 = Cross section of leaf in lower portion, X 65. 8 = Cross section of
margin in lower portion, X 250. 9 = Cross section of leaf in upper portion, X 65.
10 = Cross section of margin in upper portion, X 250. 11 = Cross section of a few
lamellae, X 250.
ells high. Vein vanishing in the apex, sharply spinose at the back,
J-^o- the leaf width. Cells above hexagonal or elliptic-hexagonal
with the longer axis transverse, .018-. 020 mm.; basal elongate-rect-
angular.
Calyptra pale, rough at apex, covering about ^ of the capsule.
Capsule cylindric, width to length about as i : 6-8, strongly arcu-
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
279
ate, brown. Teeth long, lanceolate, obtuse, median line orange.
Lid subulate, curved, one-half to once length of capsule. Pedicels
terminal, erect, flexuose, reddish-brown, 2.5-6 cm. long, 1-3 from
Fig. 3. Catharinea undulata.
1 = Dry antheridial plant; d = antheridial disk through which stem has continued,
X i. 2 = Moist plant with capsule, XL 3 = Capsule, X 5. 4 = Peristome,
X 150. 5 = Leaf, showing paired serrae, and lamellae on upper side, X 15. 6 =
Leaf-tip, X 65. 7 = Cross section of leaf, X 65. 8 = Cross section of leaf-margin
showing thickened border cells, X 250. 9 = A few lamellae in cross section, X 250.
the same stem-tip. — On clayey soil. — Juneau, Alaska, and south to
California, eastward across the continent in Canada and northern
United States; Europe; Asia.
280
FRYE
Fig. 4. Catharinea selwyni.
1 = Dry plant, showing innovation, XL 2 = Moist plant with capsule, X i.
3 = Capsule, X 5- 4 = Peristome, X 150. 5 = Leaf showing lamellae on upper
side and paired serras at margin, X 15. 6 = Leaf-tip, X 65. 7 = Cross section of
leaf, X 65. 8 = Cross section of leaf margin showing thickened border cells, X 250.
9 = Cross section of a few lamellae, X 250.
4. Catharinea selwyni (Aust.) E. G. Brit., in Bull. Torr. Bot. Club.
16: no (1889).
Atrichum selwyni Aust., in Bot. Gazette, 2: 95 (1877).
Catharinea rosulata2 Kindb., in Eur. and N. Amer. Bryin. p. 154
(1897).
2 For the reasons for including C. rosulata under C. undulata, see Bryologist, 10:
53
THE POLYTRICHACE^E OF WESTERN NORTH AMERICA 281
Named after Selwyn.3
Plants dioicous. Stems more slender than in C. undulata.
Leaves undulate when moist, with transverse rows of teeth on the
back, broader in proportion than in C. angustata, subspatulate, gen-
erally obtuse, excavated at base. Margin bordered, serrate to mid-
dle or below. Lamellae 4 — 6,9 — 13 cells high. Vein vanishing in
apex, toothed at back. Cells .025 — .050 mm.
Calyptra quite smooth.
Capsule nearly erect or subarcuate, narrowly cylindrical, dark pur-
ple, shining, width to length about as 1:6 — 8. Teeth shorter than
in Cundulata. Lid shorter rostrate than in C. undulala. — On
clayey soil and in crevices of rock. — Revelstoke, British Columbia;
Rogers Pass and Beaver Creek in Selkirk Mountains, British
Columbia; Lesser Slave Lake, Athabasca, Canada; Kootenai
County, Idaho.
OLIGOTRICHUM Lam. & DC.
Name derived from oligo = few, and tricho = hair; referring to
the almost naked calyptra.
Plants dioicous. Stems simple, 1 — 3 cm. tall, with rhizoids at base.
Leaves not undulate, i cell thick; upper leaves lanceolate to ligu-
late, when dry incurved-hooked and rarely crisped, when moist
patent to squarrose from an indistinct sheath-like base, usually with
lamellae on back toward apex; lamina on back not toothed (except
sometimes in 0. parallelum). Margin not bordered, sometimes
inflexed. Lamellae on upper side 3 — 13 (in our species), wavy from
side to side (except in 0. parallelum), with crenulate margins (except
in 0. parallelum), composed of similar smooth cells.
Calyptra with a few erect hairs, rarely smooth.
Capsule erect or inclined, symmetric, terete, oval or ovate, usually
straight, smooth, with very large 2-celled stomates. Peristome pre-
sent; teeth 32 (at least in ours), usually equal, pale throughout. Lid
readily dropping off, thinly rostrate from a conic base.
Number species in western North America, 3; total number spe-
cies, about 10.
3 Alfred Richard Cecil Selwyn, Director of the Canadian Geological Survey from
1869 to 1895; editor of the Geological and Natural History Survey of Canada, and a
large contributor to the same.
282 FRYE
THE WEST NORTH AMERICAN SPECIES, A COMPARISON AND KEY.
i. Lamellae straight; capsule with few stomates.
2. Leaf margin plane.
3. Lamellae on upper side 4-7 ; 3-6 cells high.
4. Lamellae on back low or wanting i. O. parallelum.
i. Lamellae wavy from side to side; capsule with numerous stomates.
2. Leaf margin plane.
3. Lamellae on upper side 5-7 ; 3-7 cells high.
4. Lamellae on back high 2. O. aligerum.
2. Leaf margin incurved.
3. Lamellae on upper side 10-13; 6-12 cells high.
4. Lamellae on back low or wanting 3-O. incurvum.
1. Oligotrichum parallelum (Mitt.) Kindb., in Eur. and N. Amer.
Bryin., p. 156, (1897).
Atrichum parallelum Mitt., in Journ. of Linn. Soc., 1864, p. 48, t. 8.
Atrichum leiophyllum* Kindb., in Bull. Torr. Bot. Club. 17: 275,
(1890).
Oligotrichum leiophyllum Kindb., in Eur. and N. Amer. Bryin.,
p. 156 (1897).
Probably so named because there are often tooth-like processes
parallel to the vein on the backs of the leaves.
Plants dioicous, loosely caespitose, dark green; antheridial plants
more slender, the antheridial disks cupshaped. Stems simple,
erect, 1—3 cm. tall.
Upper leaves half open, incurved when moist, slightly undulate,
ligulate-lanceolate, often marked on the back by small thin tooth-
like processes parallel to the vein; lower leaves shorter, oblong, more
obtuse. Margin plane, sharply dentate from the middle or below.
Lamellae on upper side 4 — 7, straight. Vein percurrent, sometimes
with 1—3 longitudinal dentate lamellae on back. Cells round-
hexagonal, basal cells narrow. Perichaetial leaves oblong, convolute
at base, gradually narrowed, lanceolate.
Capsule as 1:3 — 4, subcylindric, slightly arcuate, contracted
under the mouth, with few stomates. — On soil. — Kodiak Island,
Port Etches, and Douglas Island, Alaska; Vancouver Island, and
Rocky Mountain region of British Columbia; Washington.
4 See Proc. Wash. Acad. Sci., 4: 326, (1902).
Fig. 5. Oligotrichum parallelum.
1 = Antheridial plant dry; d = antheridial disk, termination of a year's growth,
X i. 2 = Moist plant with capsule, X i. 3 = Capsule with lid, X 5. 4 = Peri-
stome showing wide membrane and doubling of teeth, X 150. 5 = Leaf showing
lamellae on upper surface, X 15. 6 = Leaf showing teeth and lamellae on back, X 15
7 = Leaf-tip. X 65. 8 = Cross section of leaf showing lamellae, X 65. 9 = Cross
section of leaf margin, X 250. 10 = Cross section of a few lamellae showing smooth
and unthickened marginal cells, X 250.
284
FRYE
2. Oligotrichum aligerum Mitt., in Journ. of Linn. Soc., 1864, p.
48, t. 8.
Name derived from aliger = wing; referring to the conspicuous
lamellae on the back.
Plants dioicous, loosely caespitose, gregarious; male plants shorter,
proliferous from the center of the antheridial disk. Stems 1—3 cm.
tall, slender, radiculose at base.
Fig. 6. Oligotrichum aligerum.
1 = Antheridial plant; d = antheridial disk through which the stem has grown.
2 = Plant T7ith capsule. 3 = Capsule. 4 = Calyptra, with its few hairs. 5 = Hair
of calyptra. 6 = Peristome. 7 = Leaf, showing lamellae on upper surface. 8 =
Leaf-tip. 9 = Cross section of leaf showing lamellae. (After Sullivant).
Leaves open or spreading, oblong-lanceolate, smooth, blunt,
pointed, with lamellae on both sides. Margin plane, entire at base-
short-dentate above middle. Lamellae on upper side 5 — 7, 5 wavy
from side to side; lamellae on back 6 — 8, high, dentate. Vein per-
BBrotherus, in Engler & Prantl: Die naturlichen Pflanzenfamilien, Teil. i, Abt. 3
s. 674, shows ii in a figure.
THE POLYTRICHACE^E OF WESTERN NORTH AMERICA
current or vanishing, keeled. Cells round-quadrate, distinct. Peri-
chaetial leaves ovate, sheathing at base, erect, narrower, subulate to
apex, their cells round and pellucid.
Calyptra with a few hairs above.
Capsule long, subcylindric, slightly arcuate, contracted under the
mouth, ventricose below, with numerous stomates. — On wet clay
banks. — Rocky Mountains and coast of British Columbia;
Washington; Oregon.
3. Oligotrichum incurvum (Huds.) Lindb., in Hartm. Skand. Fl. 9
ed. 2: p. 45 (1864).
Oligotrichum hercynicum Lam. & DC. Fl. Fr. 3 ed., 2: 492 (1805).
Fig. 7. Oligotrichum incurvum.
1 = Plant with capsule, dry, XL 2 = Moist plant, XL 3 = Capsule, X 5.
4 = Peristome, X 150. 5 = Leaf showing upper side with lamellae and in-
curved margin, X 15. 6 = Leaf from back showing ridges, those on vein toothed,
X 15. 7 = Leaf- tip showing serration of margin, X 65. 8 = Cross section of leaf
showing lamellae on upper side and ridges on back, X 65. 9 = Cross section of leaf
margin X 250. 10 = A few lamellae in cross section, X 250.
286 FRYE
So named because the leaves are strongly incurved or twisted when
dry.
Plants dioicous, loosely caespitose, glaucous green, reddish-brown
when old. Stems 1 — 3 cm. high, erect, rigid, simple.
Leaves erect or spreading, lanceolate from an oblong base, more or
less acute; when dry strongly incurved and twisted but less crisped
and undulate than in Catharinea. Margin not bordered, incurved
at least above thus making leaves somewhat tubular near tip,
remotely and minutely dentate at apex but sometimes entire. Lam-
ellae on upper side 10 — 13, wavy from side to side, 6 — 12 cells high,
their margins variously notched and crested. Vein with 2—3
lamellae on back; back lamellae short, blunt, low, serrate, rather ridges
than lamellae, sometimes wanting. Cells hexagonal, rectangular at
base ; cell- walls meeting margin of leaf perpendicularly.
Calyptra with a few scattered hairs.
Capsule ovate-cylindric, erect or somewhat inclined, somewhat
irregularly plicate when dry, contracted below the mouth, with
numerous stomates. Teeth short, unequal. Lid shortly rostrate,
oblique, often falling off with the calyptra. Pedicel smooth, rather
thick, 2—4 cm. long. — On soil. — Rogers Pass, Selkirk Mountain,
British Columbia; Greenland; Europe.
COMPARISON OF VARIETY WITH TYPE.
3. O. incurvum, typical.
i. Cells wall in upper half of leaf approaching leaf margin perpendicu-
larly.
2. Cells about midway between base and tip .oio-.oi5 mm. in their
longer diameter.
3. Leaves usually remotely dentate, but sometimes entire.
4. Vein usually with 1-3 low serrate ridges or lamellae on back,
but sometimes smooth.
5. Capsule somewhat irregularly plicate.
3a. O. incurvum var. latifolium6 (C. M. & Kindb).
Oligotrichum hercynicum var. latifolium C. M. & Kindb., Mac. Cat.
VI, p. 149, (1892).
Oligotrichum integrifolium Kindb., in Revue Bryol. 1894, p. 40.
6 Name derived from latum = broad, and folium = leaf; referring to the leaves
being broader than in the type.
THE POLYTRICHACE^E OF WESTERN NORTH AMERICA
i . Cell walls in upper half of leaf approaching leaf margin diagonally.
2. Cells about midway between base and tip .020-. 030 mm. in their
longer diameter.
3. Leaves entire or nearly so.
4. Vein smooth at back.
5. Capsule distinctly plicate.
3 a
Fig. 8.
Comparison of Oligotrichum incurvum (upper figure) with Oligotrichum
incurvum var. latifolium (lower figure).
1 and la = Leaves, upper side, one narrower than the other. In 1 the margin is
usually serrate where it is rolled in, X 15. 2 and 2a = Leaf-tips, one usually with
serrate lamellae on the back, the other not, X 65. 3 and 3a = Portions of leaves show-
ing difference in size of leaf cells; also the cell walls meeting the margin perpendicu-
larly in 3 and diagonally in 3a, X 500.
On Soil — St. Lawrence Island, Bering Sea.7
7 Kindberg reports this from Rogers Pass, Selkirk Mountains, B. C-, but an examina"
tion of this material shows that in areolation, direction of marginal cell walls, and size
of leaves it is nearer to the type than to the variety. The leaf margin of O. incurvum
sometimes approaches entirety, and the back smoothness. 0. incurvum var. lati-
folium is therefore known only from St. Lawrence Island.
Proc. Wash. Acad. Sci., August, 1910.
288 FRYE
PSILOPILUM Brid.
Name derived from psilos = bare, and pilos = felt; referring to
the absence of felted or matted hairs on the calyptra.
Plants dioicous. Stems simple, from subterranean shoots.
Leaves keeled or incurved, lanceolate or ligulate, not undulate,
never toothed at back, i cell thick, when moist more or less patent
from an indistinct sheath, when dry appressed or the tips incurved.
Margin not bordered. Lamellae on upper side only, wavy from side
to side, with crenulate edges; marginal cells similar to the others.
Cells quadratic or round-hexagonal, rectangular at base.
Calyptra cucullate, naked, or at tip papillose.
Capsule usually inclined, more or less distinctly obliquely ovate,
with small mouth, laterally compressed, smooth, with large 2-celled
stomates. Peristome present (in North American species); teeth
usually unequal in size. Lid easily falling off, pointed to long and
thinly rostrate from a conic base. Pedicels single.
Number of species in western North America, i ; total number spe-
cies, about 13.
1. Psilopilum glabratum (Wahl.) Holz., in Bryologist, 5: p. 80
(1902).
Oligotrichum glabratum (Wahl.) Lindb., in Muse. Scand. p. 12 (187 9).
Psilopilum tschulschicum8 (C. M.) Par., in Index, ed. i,p. 1040.
(1897).
Psilopilum arcticum Brid., in Bryol. Univ. Vol. 2, p. 96 (1827).
Name from glabrare = to deprive of hair; referring to the smooth
calyptra.
Stem 1 — 3 cm. high.
Leaves very concave, muticous. Margin irregularly crenuiate
above. Lamellae 7 — 10, disappearing toward the base. Vein van-
ishing in the apex, smooth on the back. Cells small, long-rectangu-
lar at the base.
Calyptra very narrow, smooth.
Capsule ferruginous, black when old, ovate-gibbous. Teeth long,
some of them 2-parted, thin. Lid short, convex-conic, with short
incurved beak. Pedicel terminal, erect, brownish, .5 — 1.5 cm. long.
8 Cardot & Theriot in Proc. Wash. Acad. Sci. 4, p. 327 (1902).
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
289
—On soil. — Port Clarence, and near Nome, Alaska; St. Paul and
St. Matthew islands in Bering Sea, Alaska; Klondike River in
Yukon Territory; Labrador; Greenland; Scandinavian Peninsula;
Siberia.
Fig. 9. Psilopilum glabratum.
1 = Dry plant, XL 2 = Moist plant with capsule, X i. 3 = Capsule, X 5.
4 = Peristome, X 150. 5 = Leaf showing incurved margins and lamellae on upper
side, X 15- <5 = Leaf-tip, muticous, keeled, X 65. 7 = Cross section of leaf, X 65.
8 = Cross section of a few lamellae, X 250.
BARTRAMIOPSIS Kindb.
Bartramia is another genus of mosses, opsis = looking like; hence
looking like Bartramia.
Plants dioicous, blackish-green to brown. Stems slender, 2—8
cm. high, simple or dichotomous above, with rhizoids at base.
2QO FRYE
Leaves distant, squarrose, more or less crisped when dry, linear-
lanceolate from a hyaline sheath-like base whose edges bear each
3 — 5 long hairs, not transversely undulate, not toothed at back;
lamina 2 cells thick. Margin not bordered, densely and sharply ser-
rate. Lamellae 5 — 8, not wavy from side to side, on upper side only,
6 — 8 cells high, toothed. Vein strong, percurrent. Cells round-
hexagonal, thick-walled, .008 mm. ; sheath-cells elongated-rectangu-
lar, thin-walled.
Calyptra cucullate, naked, covering only the lid.
Capsule erect, symmetric, terete, wide mouthed, smooth, with
large 2-celled stomates. Peristome wanting. Lid conic, with long
beak. Pedicels single, 8 — 12 mm. long, reddish, flexuose when dry.
Number of species in western North America, i ; total number spe-
cies, i.
1. Bartramiopsis lescurii (James) Kindb., in Rev. Bryol. 1894,
P- 35-
Atrichum lescurii James, in Bull. Torr. Bot. Club. 6: 33 (1879).
Bartramiopsis sitkana Kindb.,9 in Rev. Bryol., 1894, p. 35.
Named after Lesquereux10
Plants laxly caespitose. Stems filiform, flexuous, laxly foliose,
long naked below.
Leaves 4 mm. long, subvaginate at base, acuminate, 2 cells thick
except near the margin where they are i cell thick, when dry very
much crisped, when moist arcuate-spreading. Margin plane, at
sheath-like base entire, with 3 — 5 hairs at edge where sheath joins
blade, further up the hairs shorten into strong teeth. Vein broad,
smooth at base. Cells of sheath hyaline, width to length about as
1:4 — 6.
Calyptra glabrous, shortly acuminate.
Capsule at first slightly ovate-cylindrical, turbinate when old, lid
long conic, long acuminate, almost equaling the capsule. Spores
9 Proc. Wash. Acad. Sci. 4; 326 (1902).
10 Leo Lesquereux, 1806-1889. A noted worker in American fossil plants and in
the mosses. With W. S. Sullivant he published "Icones Muscorum;" and with T.
P. James, "Manual of the Mosses of North America." These are today two of our
best books on North American mosses.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
291
ovate or somewhat spherical, .012 — .016 mm. — On soil. — Virgin
Bay, Orca, Douglas Island, and Wrangel, Alaska; Japan; Kam-
chatka.
Fig. 10. Bartramiopsis lescurii.
1 = Plant dry, X i. 2 = Moist plant, with capsule, X i. 3 = Moist plant, with-
out capsule, XL 4 = Capsule with lid, not quite mature and therefore slightly
shrunken, X 5. 5 = Mature capsule without lid, X 8. 6 = Leaf showing hairs at
margin and lamellae on upper side, X 15. 7 = Leaf showing narrower blade and more
curved tip, X 12. 8 = Leaf tip, upper side, showing lamellae, X 100. 9 = Leaf
tip, under side, X 100. 10 = Cross section of leaf showing 2 layers of cells except
near the margin, X 65. 11 = Cross section of a few lamellae, X 250. (Numbers
5, 7, 8, 9, after Engler & Prantl.)
POLYTRICHADELPHUS Mitt.
Polytrichum is another genus of mosses, adelphos = brother;
hence a brother to Polytrichum.
Plants dioicous, more or less robust, rigid, loosely caespitose.
Stems from subterranean rhizomes, erect or inclined, usually quite
long, densely leafy, simple or forked or tufted.
2Q2 FRYE
Upper leaves erect to patent, when dry more or less closely applied
to the stem, narrowly lanceolate to awl-shaped from a thin sheath-
like base, smooth at back; lamina i cell thick, but narrow. Margin
not bordered, slightly if at all incurved, mostly sharply toothed to
hairy above (not hairy in North American species). Lamellae on
upper side only, numerous, not wavy from side to side, entire, mar-
ginal cells mostly somewhat enlarged and somewhat oval. Vein
broad outside of sheath, excurrent as a red point. Cells of sheath
without chlorophyll, elongated-rectangular to linear, narrower
toward the margin; cells of limb iso-diametric, thick walled.
Calyptra cucullate, usually with short hairs at tip, sometimes
smooth.
Capsule inclined, oblong or ovate, 2 — 4 angled, smooth, often
semilunar in cross section, with numerous 2-celled stomates. Peri-
stome present; teeth pale, with yellowish axis. Lid conic, more or
less beaked. Pedicels single or two on a tip, elongated, thick.
Number of species in western North America, i ; total number of
species, about 18.
1 , Polytrichadelphus lyallii Mitt. , in Journ. Linn. Soc. 1 864, p. 49.
Oligotrichum lyallii Lindb., in Act. Soc. pr. Fauna et Fl. Fenn, 1868,
p. 102.
Polytrichum angustidens Lindb.11 in Bot. Centralb. Vol. 84.
Named after Lyall.12
Plants robust, dirty yellow to brownish green. Stems fastigiately
branching from the middle, sometimes simple, naked at the base,
densely foliate above.
11 Prof. J. M. Holzinger, of Winona, Minnesota, kindly sent some Polytrichum angus-
tidens Lindb., co-type material, No. 1121 of the collection by Sandberg in northern
Idaho. A comparison of this with Polytrichadelphus lyallii Mitt, indicates that they
are the same. The leaf characteristics even down to the marginal cells of the lamellae
agree. The number of lamellae is 35-45. The capsules sent were young and shrunken;
the angles cannot therefore be made out with certainty. Lindberg says, "capsules
acutely 4-angled," but if his material was likewise young and shrunken, one could
easily surmise an error here. The other capsule characteristics agree with Polytricha-
delphus lyallii. The calyptra would at once distinguis h Polytrichum from Polytricha-
delphus, but it is wanting in the writer's material; since Lindberg omits it in his
description, one surmises it was wanting in his as well. More evidence is necessary
to convince the writer that Polytrichum angustidens is not Polytrichadelphus lyallii.
12 David Lyall, surgeon and botanist attached to international survey.
Fig. 11. Polytrichadelphus lyallii.
1 = Plant moist, with capsules, X i. 2 = Plant dry, Xi. 3 = Capsule, showing
double fold. 4 = Capsule, X 5. 5 = Cross section of capsule near base, X 10.
6 = Cross section of capsule near mouth, X 10. 7 = Calyptra. 8 = Hair of calyp-
tra. 9 = Peristome. 10 = Leaf. 11 = Leaf, X 15. 12 = Leaf tip, X 65. 13 =
Cross section of leaf. 14 = Cross section of a few lamellae, X 250. (Nos. 3, 7, 8, 9,
10, 13, after Sullivant.)
2Q4 FRYE
Leaves oblong, clasping at base, narrowly lanceolate above, con-
vex and smooth on back. Margin incurved, distantly serrate from
the middle up. Lamellae 35 — 45, 3 — 7 cells high; marginal cells
rough, oval, higher than wide. Inner perichaetial leaves with long
convolute base and short acumen.
Calyptra fugacious, with few appressed hairs, split on one side, not
curved, slightly twisted.
Capsule slightly inclined, cylindric-oblong, ventricose below,
bluntly 2 — 4 angled,13 plicate-rugose at base when empty. Teeth
64. Lid broadly conic, subulate-rostrate. Pedicel long, flexuous.
— On soil. — In mountains from British Columbia southward to
Colorado, Nevada, and California.
POGONATUM P. Beauv.
Name derived from pogon = beard; referring to the hairy calyp-
tra.
Plants dioicous (in species included), loosely caespitose, olive to
dark green. Stems erect, simple or branched above.
Leaves when dry crisped, appressed or spreading more or less, lan-
ceolate to linear-lanceolate, usually with a sheath-like base, sheath
and lamina i cell thick; lamina smooth at back. Margin not
incurved, usually sharply serrate, not bordered. Lamellae mostly
very numerous (at least 35 in the included species), not at all or very
little wavy from side to side, entire, rarely few to wanting. Vein
toothed at back toward apex, sometimes smooth, wide, many cells
thick. Cells of lamina small, thick walled, round-hexagonal; cells of
sheath rectangular to linear.
Calyptra with long smooth hairs at tip which usually form a felted
mat covering the whole surface.
Capsule erect or inclined, straight, sometimes slightly curved,
terete, sometimes slightly ribbed, cells of epidermis usually mamil-
lose; without stomates, except in P. alpinum] hypophysis wanting or
13 This species is described in Lesquereux & James' Manual as having the capsule
biplicate and semilunar in cross section. There are however two other faint angles
extending part way up from the base, making it 4-angled at least below. Brotherus'
description of the capsule of the genus (Engler & Prantl. Nat. Pflanzenf. Teil i, Abt.
3, s. 682) as " 2-kantig, in Querschnitt halbmondf ormig " does not hold as shown by the
cross sections of the capsule in the plate.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 295
indistinct; Peristome present; teeth 32 (in the included species), in
P. alpinum doubled so one might count 64. Lid hemispheric, more
or less long-beaked. Pedicels single, rarely several at one tip, more
or less elongated, purple.
Number of species in western North America, 4; total number spe-
cies, about 133.
THE WEST NORTH AMERICAN SPECIES, A COMPARISON AND KEY.
i. Leaves very much crisped when dry.
2. Marginal cells of lamellae smooth, not thickened.
3. Teeth 32.
4. Capsule without stomates.
5. Capsule papillose.
6. Marginal cells of lamellae differing little in size and form
from the others i. P. contortum.
i. Leaves hardly or not at all crisped when dry.
2. Marginal cells of lamellae papillose or rough, thickened.
3. Teeth 64, or 32 double ones.
4. Capsule with stomates. '
5. Capsule not papillose.
6. Marginal cells of lamellae ovate, width to length as i :
J-2, larger than the others 4. P. alpinum.
3. Teeth 32.
4. Capsule without stomates.
5. Capsule papillose.
6. Marginal cells of lamellae oval or flat- topped, width to
length as 1^-2:1, larger than the others.
2. P. capillare.
6. Marginal cells of lamellae round, about the same size as
the others j. P. urnigerum.
1, Pogonatum contortum (Menz.) Lesq., in Mem. Calif. Acad. i,
p. 27.
Pogonatum erythrodontium Kindb., in Mac. Cat. p. 150 (1892).
Pogonatum atrovirens Mitt.,14 in Journ. Linn. Soc. 1864, p. 49.
14 An examination of P. atrovirens Mitt, shows it to be P. contortum. Type material
from the Mitten Herbarium, now owned by the New York Botanical Garden, and also
material collected by Macoun at Hastings, Burrard Inlet, near Vancouver, British
Columbia, was compared with authentic P. contortum. The capsules are papillose
as in P. contortum. The basilar areolation is the same in leaves taken from correspond-
ing parts of the stem. The leaves in both vary in the size of the sheath, those near
the base of the stem having larger sheaths than those near the tip. Since these con-
stitute the characteristics upon which P. atrovirens Mitt, is founded, it reduces to
P. contortum.
Fig. 12. Pogonatum contortum.
1 and 2 = Dry antheridial plants; d = antheridial disks through which stems have
grown, X i. 3 and 4 = Moist plants with capsules; stems have continued beyond point
where pedicel arises, XL 5 and 6 = Capsules, X 5. 7 = Immature capsule covered
by calyptra, X 5. 8 = Peristome, X 150. 9 and 10 = Leaves, showing lamellae
on upper side, X 15. 11 = Leaf tip, X 65. 12 and 13 = Cross sections of leaves,
X 65. 14 = Cross section of leaf margin, X 250. 15 and 16 = Cross sections of
lamellae, X 250.
THE POLYTRICHACE^E OF WESTERN NORTH AMERICA 297
So named because the leaves are strongly twisted or contorted
when dry.
Plants large, gregarious or loosely caespitose, glaucous green above,
brown below. Stems simple, or with an innovation from under the
perichetium, loosely and irregularly foliate its whole length.
Leaves erect, open, twisted and crisped when dry, linear-lanceo-
late, usually longer upwards on the stem, acute, sheath scarcely
broader than blade. Margin sharply serrate to the base. Lamellae
20 — 40. Marginal cells of the lamellae oval, smooth, not very
much larger than the others. Vein percurrent, sparingly dentate on
the back. Perichaetial leaves similar to the foliage leaves.
Calyptra covering the whole capsule.
Capsule ovate to obovate or cylindric, erect or somewhat curved,
papillose, when dry slightly constricted under the mouth, without
stomates. Teeth 32. Lid convex, rostellate. Pedicel long, flexu-
ous. — On soil, usually clay. — Along the coast from the Alaska Penin-
sula to California; Rocky Mountains, at least of British Columbia.
2. Pogonatum capillare (Rich.) Brid., in Bryol. Univ. II, p. 127,
(1827).
Pogonatum dentatum Brid., in Bryol. Univ. II, pp. 122 and 744,
(1827).
Pogonatum capillare var. dentatum Lindb.,15 in Act. Soc. sc. Fenn.
1872, p. 266.
Name derived from capillaris = hairy; probably referring to the
hairy calyptra, so common in this family.
Plants 2.5 cm. high or less, gregarious or loosely caespitose, glauc-
ous green; male plants smaller. Stems slender, mostly simple,
loosely foliate, with rhizoids at base.
15 Cardot and Theriot, in "Mosses of Alaska, " Proc. Wash. Acad. Sci. 4: 327 (1902)
say "Pogonatum dentatum (Menz.) Brid. is but a western race of P. capillare, charac-
terized by having slenderer stems than those of the type, and by its pedicel which is
usually not so flexuous. " This hardly seems sufficient difference for a variety.
Cloudy weather and wet soil cause stems to grow longer and more slender. The
northwest coast of North America is characteristically damp and cloudy. Potatoes
growing in a dark damp cellar are not called new varieties. Further the difference
is not constant. The marginal cells of the lamellae vary a great deal, so this dis-
tinction, shown in the figures in Sullivant's Icones Muscorum, does not hold.
Fig. 13. Pogpnatum capillare.
1 = Moist plants, with capsules, XL 2 = Dry antheridial plant; d = antheridial
disks, X i. 3 and 4 = Immature capsules covered by calyptras, X 5. 5 and 6 =
Capsules with lid. X 5. 7 = Peristome, X 150. 8 = Double tooth of peristome, X
150. 9 and 10 = Leaves showing sheath at base and lamellae on upperside, X 15.
11 and 12 = Leaf tip, X 65. 13 and 14 = Cross sections showing lamellae and leaf
margins, X 65. 15\nd 16 = Cross sections of a few lamellae, X 250.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
Leaves not crisped when dry, gradually longer upwards; lower
leaves distant, small, appressed; upper leaves large, linear from a
short sheathing base. Margin sharply serrate. Lamellae 45 — 55;
marginal cells of lamellae much larger, papillose, rectangular or oval,
wider than long as 1^ — 2:1.
Calyptra hairy, hardly covering capsule to base.
Capsule oblong-cylindric, erect, papillose, thin, without stomates.
Teeth 32. Lid hemispheric, abruptly straight-beaked. Pedicel
slender, flexuous, long.— On soil.— St. Paul Island, Bering Sea;
from the Alaska Peninsula along the coast to southern Alaska;
Rocky Mountains of British Columbia and the United States;
Portland, Oregon; Adirondack Mountains; White Mountains; New-
foundland; Miquelon Island; Greenland; Scandinavian Peninsula;
Siberia.
3. Pogonatum urnigerum (L.) Beauv., in Prodr. p. 84 (1805).
Polytrichum urnigerum L., in Sp. PI. II, p. 1109. n. 3 (1753).
Name derived from urna = urn, and gerere = to bear; probably
referring to its erect, wide-mouthed capsule.
Plants erect, 2.5 — 7.5 cm- high> dark green, brown below. Stems
rigid, sometimes forked at tip.
Upper leaves lanceolate from a pale sheathing base, crowded, at
apex acute to acuminate; when moist patulose; when dry rigid, not
crisped, appressed, erect. Margin plane or erect, sharply toothed
nearly to sheath. Lamellae 40 — 50, 4 — 6 cells high; marginal cells
not much larger, rounded, thickened, papillose, yellowish-green.
Vein slightly excurrent or vanishing, sharply spinose at back. Cells
quadrate-hexagonal or rounded.
Calyptra longer than the capsule.
Capsule erect or nearly so, symmetric, wide-mouthed, yellowish-
brown to brown, without stomates, very papillose. Teeth 32, red-
dish. Beak of lid almost straight, subulate. Pedicel 2.5—4 cm.
long, slender, pale reddish. — On soil and soil-covered rocks. —
Coast of Alaska from Disenchantment Bay near mouth of Yukon
River to the vicinity of Juneau; Rogers Pass, Selkirk Mountains,
British Columbia; Cathlamet, Washington; Portland, Oregon;
Europe; Asia.
3oo
FRYE
Fig. 14. Pogonatum urnigerum.
1 = Dry antheridial plant; d = old antheridial disk through which young shoot has
grown, X i. 2 = Moist plant with capsule, XL 3 = Calyptra covering capsule,
immature, XL 4 = Capsule, X 5. 5 = Peristome, X 150. 6 and 7 = Leaves
showing lamellae on upper side, X 15. 8 = Leaf tip, X 65. 9 = Cross section of
leaf, X 65. 10 = Cross section of a few lamellae, showing rounded, thick-walled
papillose, marginal cells, X 250.
THE POLYTRICHACE^E OF WESTERN NORTH AMERICA 301
4. Pogonatum alpinum (L.) Roehl., in Ann. Wett. Gesells. Ill, p.
226 (1812).
Polytrichum alpinum L., in Sp. pi. II, p. 1109, n. 2 (1753).
Probably so named because it is found in mountain (alpine) regions
in Europe.
Plants loosely or densely tufted, tall, decumbent at base. Stems
much branched, rarely simple.
Leaves dull green, narrowly acuminate, not crisped when dry.
Margin serrate. Lamellae 30-40, 5-8 cells high; marginal cells
slightly larger than the others, ovate to ovate-conic, longer than
wide, thickened, papillose or rough, yellowish.
Calyptra shorter than the capsule.
Capsule inclined, arcuate, subglobose to elongate-cylindric and
curved, terete, narrower at mouth than below, smooth, with stom-
ates, greenish-brown, black and rugose when old; hypophysis in-
distinct or wanting. Peristome present; teeth 64, or 32 double
ones, short, irregular. Beak of lid curved, long, subulate. Pedi-
cel long, flexuous. — On soil. — From Kotzebue Sound north of
Bering Strait in Alaska southward to Washington and Idaho;
White Mountains of New Hampshire; Europe; Asia; Australia.
COMPARISON OF VARIETIES WITH TYPE.
4a. P. alpinum var. simplex16 Sch., in Coroll. p. 91, (1856).
i. Leaves about 5 mm. long. "
2. Leaf-sheath to blade about as 1 12 J.
3. Plant about 2 cm. tall or shorter.
4. Capsule i . 8-2 . 3 mm. long.
5. Width of capsule to length as i : i-J— 2j
6. Stems simple.
7. Pedicel i|-2 cm. long.
On soil and rocks. — Port Clarence, Alaska; Colorado; Copper
Mountains and Gold range in British America.18
16 So named on account of its unbranched stems.
17 It should be borne in mind that varietal differences are not so constant as those
characterizing species; varieties grade into each other more. Therefore, while these
numbered statements constitute a description and comparison of the varieties, con-
siderable latitude must be permitted for variations.
18 An examination of Macoun's No. 427 leads to the conclusion that this is var.
simplex instead of var. septentrionale.
302
FRYE
Fig. 15. Pogonatum alpinum.
1 = Plant dry, X i. 2 = Moist plant with capsule, X i. 3 = Capsule, X 5
4 = Peristome, X 150. 5 = Leaf with sheathing base and lamellae on upper side,
X 15. 6 = Leaf tip, X 65. 7 = Cross section of leaf showing lamellae and margin,
X 65. 8 = Cross section of a few lamellae showing thickened and papillose marginal
cells, X 250.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 303
4b. P, alpinum var. brevifolium19 Brid., in Sch. Syn. I. ed. p.
441, (1860).
Polytrichum alpinum var. brevifolium Muell., in Syn. I, p. 210, (1849).
i. Leaves about 7-8 mm. long.
2. Leaf-sheath to blade about .as 1:3-3?.
3. Plant about 4 cm. tall or shorter.
4. Capsule 3-4 mm. long.
5. Width of capsule to length as i :i£-2i.
6. Stems with few* and short branches.
7. Pedicel 2^-3^. cm. long,
a. Capsule with distinct neck.
On rocks. — Islands of Bering Sea; Mt. Dana, California; Hudson
Strait; Greenland; mountains of Europe; Siberia.
4c. P. alpinum var. septentrionale20 (Sw.) Brid., in Sch. Syn.,
i ed., p. 44 1 (1860).
P. alpinum var. microdontium?1 Kindb., in Mac. Cat. Vol. 6, p.
152, (1892).
i. Leaves about 7-8 mm. long.
2. Leaf-sheath to blade about as 1 13-34.
3. Plants about 5 cm. tall or shorter.
4. Capsule 3-4 mm. long.
5. Width of capsule to length as i :iJ-2^.
6. Stems simple.
7. Pedicel 1^-2 cm. long,
a. Capsule without distinct neck.
On soil and rocks. — Islands of Bering Sea; Kodiak Island,
Alaska; mouth of Skeena River and Rocky Mountains, British
Columbia; Europe.
4d. P. alpinum var. arcticum22 (Sw.) Brid., in Sch. Syn. i ed*
p. 441 (1860).
Polytrichum sylvaticum Menz., in Trans. Linn. Soc. 4. p. 83, n.
19 (1798).
Polytrichum alpinum var. arclicum Wahl., in Fl. Lapp., p. 346
(1812).
19 Breve = short, folium = leaf; because its leaves are shorter than those of nearly
all other varieties.
20 Septentrionale = pertaining to the north; referring to the northern habitat of
this variety.
21 Examination of Alaskan collections leads to the conclusion that Cardot and Th£riot
are right in concluding that var. microdontium does not separate from var. septentrionale-
22 So named on account of its arctic habitat.
Proc. Wash. Acad. Sci., August, 1910.
304 FRYE
i. Leaves about 7-8 mm. long.
2. Leaf-sheath to blade about as 1:3-4.
3. Plants about 5 cm. tall or shorter.
4. Capsule 4-5 mm. long.
5. Width of capsule to length as 1 13-5.
6. Stems simple or with few branches.
7. Pedicel 2-3 cm. long.
On soil and rocks. — Egg Island, Disenchantment Bay, Alaska;
Stewart Island and Mt. Rainier, Washington; northern Europe.
4. P. alpinum, typical.
i. Leaves about 9-11 mm. long.
2. Leaf-sheath to blades about as 1:3-5.
3. Plants about 10 cm. tall or shorter.
4. Capsule 4-5 mm. long.
5. Width of capsule to length as 1 12-3.
6. Stem much branched.
7. Pedicel 2^-3^ cm. long.
4e. P. alpinum var. macounii23 (Kindb.) C. & Ther. in Proc. Wash.
Acad. Sci. 4, p. 328 (1902).
P. macounii Kindb., in Bull. Torr. Bot. Club, 16, p. 96 (1889).
i. Leaves about 12-14 mm- l°ng-
2. Leaf-sheath to blade about as i :3~5.
3. Plants about 15 cm. tall or shorter.
4. Capsule 5-6 mm. long.
5. Width of capsule to length as i :3~4.
6. Stems simple, rarely branched.
7. Pedicel 4-7 cm. long.
On soil. — From the Alaska Peninsula southward along the coast
to Washington, and eastward across British Columbia to the Rocky
Mountains.
POLYTRICHUM Dill
Name derived from poly = many, and tricho = hair; referring
to the hairiness of the calyptra.
Plants dioicous, loosely to densely caespitose. Stems rigid, from
subterranean rhizomes, erect or nearly so, densely leafy, simple,
rarely forked or much branched at the tip.
Leaves erect when dry, from a sheath-like base, lanceolate to
awl-shaped, more than i cell thick except at margins, sheath i cell
23 Named after John Macoun, naturalist of the Canadian Geological Survey.
THE POLYTRICHACE^: OF WESTERN NORTH AMERICA 305
typical.
macoumi.
Fig. 16. Pogonatum alpinum, and variety.
1 = Plants moist, X i. 2 = Dry plants, X i. 3 = Capsules, X 5. 4 = Leaves,
X 15-
306
FRYE
arcticum.
brevifolium. simplex.
septentrionale.
Fig. 17. Pogonatum alpinum, varieties.
1 = Plants moist, X i. 2 = Plants dry, XL 3 = Capsules, X 5- 4 = Leaves,
Xis-
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 307
thick; lamina smooth at back. Margin plane or broadly incurved,
not bordered, usually with large teeth. Lamellae on upper side only,
very numerous, erect, high, not wavy from side to side, entire or
nearly so. Vein often toothed at back near apex. Cells of sheath
without chlorophyll, elongated-rectangular to linear, toward the
edge longer than along the vein; cells of lamina small, thick
walled, quadratic or hexagonal, at base often broader than long,
often in rows at the margin.
Calyptra matted with very long white to brown hairs which cover
the whole capsule.
Capsule erect when young, later inclined, finally often horizon-
tal, 4-6 angled or terete, cubical to oblong; hypophysis hemispheric
and grading into capsule, or discoid and deeply constricted from
capsule, with large i-celled stomates. Peristome present; teeth 64.
Lid large, plane to conic, beaked. Pedicels single, long, rigid, yel-
lowish-red to purple, twisted to the right above when dry.
Number of species in western North America, 12; total number
species, about 104.
THE WEST NORTH AMERICAN SPECIES, — A COMPARISON AND KEY.
i. Leaf margin serrate to sheath or nearly so.
2. Margin plane or erect.
3. Leaves sharply pointed, the point composed mostly of the ex-
current vein.
4. Leaf point red, dentate, about i mm. long.
5. Marginal cells of lamellae like the rest, sometimes slightly
flattened or elongated, not thickened, width to height
about as i: i.
6. Capsule about as i :i£, obscurely 5-6 angled.
7. Plants 2^-10 cm. long.
a. Cells near middle of sheath and § distance from
margin to vein about 3-4 times as long as wide;
leaf blade about middle i cell thick for 4-8 cells
from margin i. P. gracile.
6. Capsule about as i :if, sharply 4-6 angled.
7. Plants 7-20 cm. high.'
a. Cells near middle of sheath and § distance from
margin to vein about 6-10 times as long as wide;
leaf blade about middle i cell thick for 2-3 cells
from margin 2. P. attenuatum.
5. Marginal cells of lamellae oval or flattened, thickened, width
to height about as 1^-2 :i.
6. Capsule about as i : 2-2^, sharply 4-6 angled.
7. Plants 3-6 cm. high j. P. ohioense.
308 FRYE
5. Marginal cells of lamellae notched in cross section.
6. Capsule about as mJ-i-J, sharply 4-angled.
7. Plants 5-45 cm. high 5. P. commune.
4. Leaf point red, smooth, £ mm. or shorter.
5. Marginal cells of lamellae oval or flattened in cross section,
not thickened, width to height about as 2: i.
6. Capsule unknown.
7. Plants 4-8 cm. high 4. P. inconstans.
i. Leaf margin serrate from the middle up.
2. Margin plane below, erect or somewhat incurved above.
3. Leaves sharply pointed, the point composed mostly of the excur-
rent vein.
4. Leaf point red, dentate, i mm. long or shorter.
5. Marginal cells of lamellae notched in cross section.
6. Capsule unknown.
7. Plant 4-8 cm. high 6. P. jensenii.
i. Leaf margin entire.
2. Margin plane or erect.
3. Leaves sharply pointed, the vein excurrent as an arista.
4. Arista red, smooth, i mm. long or shorter.
5. Marginal cells of lamellae notched in cross section.
6. Capsule unknown.
7. Plants 5-8 cm. high 7. P. yukonense.
2. Margin plainly incurved.
3. Leaves blunt, cucullate.
4. Arista or sharp point none.
5. Marginal cells of lamellae ovate, thickened, width to height
about as i 12.
6. Capsule about as i : ij, bluntly 6- angled.
7. Plants 2.5-10 cm. high 8. P. sexangulare.
3. Leaves sharply pointed, the vein excurrent as an arista.
4. Arista red, rarely somewhat colorless at tip, rough, ^-ij mm.
long.
5. Marginal cells of lamellae ovate or flask-shaped.
6. Capsule as i : ij-ij, sharply 4- angled, 3-5 mm. long.
7. Plants 2-10 cm. high.
b. Stem without^ dense covering of rhizoids.
9. P. juniperinum.
6. Capsule about as i nj, sharply 4-angled, 2-3mm. long.
7. Stems mostly with dense covering of rhizoids.
10. P. strictum.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 309
4. Arista hyaline, rough, i^ mm. or shorter.
5. Marginal cells of lamellae ovate or flask-shaped, thickened,
width to height about as 1 11^-2.
6. Capsule about as i:i-ij, sharply 4-angled.
7. Plants 3-12 cm. high,
c. Stems mostly branched n. P. hyperboreum.
4. Arista hyaline, rough, 3 mm. or shorter.
5. Marginal cells of lamellae ovate or flask-shaped, thickened,
width to height about as i:iJ-2.
6. Capsule about as i:ij-ij, sharply 4-angled.
7. Plants 2 . 5-4 cm. high,
c. Stems simple 12. P. piliferum.
1. Polytrichum gracile Dicks., in MSS.
Name derived from gracilis = slender; probably referring to the
rather tall stem, naked below.
Plants tufted, dark green 2.5-10 cm. high. Stems matted to-
gether below with whitish tomentum, simple or slightly divided,
leafless below.
Leaves erecto-patent, somewhat flexuous or patent when dry, 8~io
mm. long, the limb lanceolate, gradually narrowed to a short acumen,
limb usually about 4-6 cells wide in the middle of the leaf. Margin
erect, sharply serrate, variable in width. Lamellae about 40, 4-6
cells high; marginal cells about equalling the others, not papillose,
rounded or a little higher than wide; vein toothed above at back,
excurrent; point short, red, dentate. Cells of limb .oi5-.oi8 mm.,
round-quadrate to transversely elliptical; cells of sheathing base
thin, rectangular, in middle of wing length to width about as i : 3-4.
Perichaetial leaves long-sheathing.
Calyptra hardly covering the capsule.
Capsule short, broadly ovate, inflated, erect, horizontal when
dry, with 5-6 obtuse and often obscure angles, smooth, narrowed
at mouth; hypophysis rather indistinct, hardly constricted above.
Teeth often confluent and unequal. Lid large with a rather long
beak. Spores .oi8-.O22 mm. Pedicel 4-5 cm. long, thin, flexuous.
—On soil. — Kotzebue Sound north of Bering Strait, and Kodiak
Island, Alaska; below White Horse Rapids, Yukon region, western
British America; Rocky Mountain region of British Columbia;
Eastern United States and British America.
310
FRYE
Fig. 18. Polytrichum gracile.
1 = Moist plant with capsule, XL 2 = Dry plant, XL 3 = Capsule, X 5.
4 = Peristome, X 150. 5 = Leaf showing lamellae on upper side, X 15. 6 = Leaf
tip, X 65. 7 = Cross section of leaf showing lamellae, X 65. 8 = Cross section of
a few lamellae showing smooth marginal cells, X 250.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
2. Polytrichum attenuatum Menz., in Trans. Linn. Soc. (1798),
p. 72.
Polytrichum conorhynchum Kindb., in Mac. Cat., p. 164 (1892).
Polylrichum formosum Hedw., in Sp. M., p. 92 (1801).
Name derived from attenuatus = drawn out; referring to the
long narrow, simple form of the plant often found.
Plants loosely tufted, dark green, 7-20 cm. high. Stems erect or
ascending, tomentose at base, simple or forked.
Fig. 19. Polytrichum attenuatum.
1 = Moist plant with capsule, XL 2 = Dry plant, X i". 3 = Capsule, X 5.
4 = Peristome, X 150. 5 = Leaf showing lamellae on upper side, X 15. 6 = Leaf
tip, X 65. 7 = Cross section of leaf showing lamellce ,X 65. 8 and 9 = Cross sections
of a few lamellae showing height and the marginal cells, X 250.
312 FRYE
Upper leaves spreading, loosely incumbent when dry, similar to
those of P. gracile but larger, 10-15 mm. long. Margin erect,
sharply serrate, very narrow, of 2-3 rows of smaller cells (.010-
.012 mm.). Lamellae about 50-70, very low, 3-5 cells high; mar-
ginal ceils smooth, round, equal or slightly larger man tne otners ana
sometimes a little longer than broad. Vein excurrent; point red,
short, dentate. Cells of leaf-base longer and narrower than in P.
gracile, very narrow at margin, near middle toward vein width to
length about as i :6-io. Perichaetial leaves with long sheaths,
longly acuminate.
Calyptra covering capsule.
Capsule erect or inclined, cernuous when dry, oblong, with 5-6
(rarely 4) acute angles, yellow-green, fawn-colored when ripe, wide-
mouthed, cells of surface turgidly mamillose, smooth; hypophysis
small, distinct, constricted above. Lid large, with rather long
beak. Spores .010 - .014 mm. Pedicel 5-6 cm. long, stout, rigid,
orange below and paler above. — On soil and rocks. — From Alaska
Peninsula southward along the coast, Ravelstoke and Selkirk
Mountains in British Columbia; Oregon; Pelee Point on Lake Erie
in eastern Canada; Miquelon Island; Europe; Japan.
3. Polytrichum ohioense R. & C., in Rev. Bryol., 1885,
p. ii.
Name derived from Ohio in which state it was first found.
Plants erect, 3-6 cm. high. Stems simple or bipartite.
Leaves spreading when moist, erect-flexuous when dry, cuspidate;
linear-acuminate from a sheathing base. Margin plane or erect »
serrate. Lamellae about 50, 5-7 cells high; marginal cens mucn
larger than the others, broader than long as 1^-2 : i, slightly con-
vex or plane. Vein excurrent in a red dentate point. Perichaetial
leaves longer, with a longer hyaline base.
Capsule erect, finally horizontal, acutely 4-5 angled, rarely 6
angled, smooth, rather narrow towards the base, 2-2^ mm. wide,
5-7 mm. long; hypophysis very small or indistinct. Lid conic-
acuminate, red at margin. Pedicel 4-6 cm. long, reddish below,
pale above.24 — On soil. — Ravelstoke, British Columbia; eastern
British Columbia; northeastern United States.
24 Examination of material from Alaska, eastern United States and Europe shows
that P. ohioense cannot be distinguished from P. attenuatum by the form of the capsule
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 313
Fig. 20. Polytrichum ohioense.
1 = Moist plant with capsule, XL 2 = Dry plant, X i. 3 = Capsule, X 5- 4 =
Immature capsule with calyptra, X 5. 5 = Peristome, X 150. 6 = Leaf showing
lamellae on upper side, X 15. 7 = Leaf tip, X 65. 8 = Cross section of leaf showing
lamellae, X 65. 9 = Cross section of a few lamellae showing usual form of marginal
cells, X 250 . 10 = Cross section of a few lamellae just above sheath, X 250.
and distinctness of the hypophysis. However, all the specimens examined could at
once be distinguished by the marginal cells of the lamellae. Occassionally in P. ohioense
they are somewhat depressed in the center approaching forms of P. Commune but the
elongated capsule clearly separates it from the last. P. ohioense tends to have fewer
lamellae than P. attenuatum; of the specimens examined none of the former had over
45, while none of the latter had less than 45.
314
FRYE
4. Polytrichum inconstans Hagen, in Nyt. Mag. Naturvid.
1900, p. 339.
Inconstans = inconstant; just how this applies to this species the
writer does not know.
Plants 8 cm. or less. Stems rather uniformly leaved above, with
a felt of dark red rhizoids below.
Leaves somewhat distant, irregularly spreading, 6 mm. long or
less, lanceolate; lamina under lamellae 2 cells thick, between lamellae
Fig. 21. Polytrichum inconstans.
1 = Moist antheridial plant; d = antheridial disk, X i. 2 = Dry plant, X i.
3 = Leaf showing lamellae on upper side, X 15. 4 = Leaf tip, X 65. 5 = Cross
section of part of leaf showing margin, height of lamellae, and form of marginal cells.
(No. 5 after Williams).
and margin i cell thick. Margins plane below, erect or somewhat
incurved above, minutely serrulate. Lamellae 24 or less, the median
ones 5-9 cells high; marginal cells sometimes 2 side by side, not
papillose, oblong, wider than long (about as 2 : i), scarcely or not
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 315
grooved. Vein scarcely or shortly excurrent, the point smooth.
Cells in limb between lamellae and margin mostly .012 by .016 mm.
to .016 by .020 mm.; cell walls on back of leaf not thickened.
Otherwise unknown. — Lake Lindeman, Yukon region of British
America; Norway; Iceland.
5. Polytrichum commune L., in Sp. PL II, p. 1109, (1753).
Commune = common. It is quite common in Europe where
Linnaeus found it.
Plants very tall, 5-45 cm. high, in large loose cushions, deep green
tomentose at base; male plants shorter, with longer leaves, repeat-
edly proliferous from the center of the antheridial disk or cup.
Stems simple, rarely forked, flexuose, wiry.
Upper leaves very long, squarrose or recurved when moist, erect
and appressed when dry, with the apex flexuose, rather laxly ar-
ranged, very long, suddenly narrowed from an oblong sheathing
base; limb lanceolate, gradually narrowing from its base upward
to a sharp dentate acumen. Margin densely and sharply serrate
to the base of the limb, very narrow, of about i row of small cells.
Lamellae about 60, low, thickened at the upper border and chan-
nelled, 4-6 cells high; marginal cells broader than the others, de-
pressed in the center and bicuspidate in section. Cells of leaf-
base and of margin similar to those of P. attenuatum. Perichaetial
leaves more distinct than in P. attenuatum, long-sheathing membran-
eous, without lamellae, ending in an arista formed by the excur-
rent vein.
Calyptra large, golden-brown, descending below the capsule.
Capsule at first erect, when ripe reddish-brown and horizontal,
4-angled, with the two lateral angles usually larger and more prom-
inent than the upper and lower, slightly flattened, shortly rectangu-
lar or almost cubic ; hypophysis discoid, very distinct, narrowly con-
stricted above; surface cells each with a high conical papilla bear-
ing a small narrowly-elliptical or slit-shaped pore at the apex. Lid
with short rostellate beak. Pedicel 6 - 10 cm. long, stout. — On soil.
— Between Cook Inlet and Tanana River, Kodiak Island, and Sitka,
Alaska; Dawson and Lake Lindeman in Yukon region of British
America; Washington; Oregon; Eastern United States and British
America; Europe; Asia; Australia; South America.
Fig. 22. Polytrichum commune.
1 = Moist plant with capsule, X i. 2 = Antheridial plant dry; d = antheridial
disk, X i. 3 = Immature capsule with calyptra, X 5- 4 = Capsule, X 5. 5 =
Peristome, X 150. 6 = Leaf showing lamellae on upper side, X 15. 7 = Leaf tip, X
65. 8 = Cross section of leaf, X 65. 9 = Cross section of a few lameLae showing
notched marginal cells, X 250.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 317
COMPARISON OF VARIETIES WITH TYPE.
5. P. commune, typical.
i. Lamellae 4-6 cells high.
2. Leaves appressed when dry.
3. Inner perichaetial leaves not exceeding the foliage leaves.
4. Width of capsule to length about as i :ij-i^.
5. Beak of lid slightly curved.
6. Pores of papillae of capsule wall from elliptical to slit-like.
5a. P. commune var. uliginosum Hueb., in Muscol. Germ. p. 535
(1833)-
i. Lamellae 4-6 cells high.
2. Leaves spreading-recurved when dry.
3. Inner perichaetial leaves not exceeding the foliage leaves.
4. Width of capsule to length about as i :ij-i J.
5. Beak of lid slightly curved.
6. Pores of papillae of capsule wall from almost rounded to
slit-like.
On soil. — White Mountains, New Hampshire; Washington.
5b. P. commune var. perigoniale25 (Michx.) B. & S., in Bryol.
Eur. fasc. 21-24, iv, i844.
i. Lamellae 6-9 cells high.
2. Leaves appressed when dry.
3. Inner perichaetial leaves very long, exceeding the foliage leaves.
4. Width of capsule to length about as i :ii-ii.
5. Beak of lid straight.
6. Pores of papillae of capsule from rounded to slit-like.
On soil. — McLeod Lake, British Columbia; Pass Creek, near
Sproat, British Columbia; Ontario; Europe; Australia.
5c. P. commune var. minus26 Weis, in PL Crypt. Goett. p. 171
(1770).
P. commune var. canadense27 Kindb., in Mac. Cat. p. 156 (1892).
i. Leaves 6-9 cells high.
2. Leaves appressed when dry.
3. Inner perichaetial leaves not exceeding the foliage leaves.
4. Width of capsule to length about as i :i-ii.
5. Beak of lid slightly curved.
6. Pores of papillae of capsule wall from elliptical to slit-like.
25 Peri = around, gone = female plant organ; referring to the perichaetial leaves
which are very long in this species.
26 Minus = small. This is one of the smaller varieties of the species.
27 The var. minus does not seem to differ from the var. canadense by any well
denned characteristics.
typical.
minus.
uliginosum. perigoniale.
Fig. 23. Polytrichum commune and varieties.
1 = Dry plants, X i. (The typical plant often has more erect leaves.) 2 = Tip
of plant showing perichaetial leaves and capsule, XL 3 = Capsules, X 5- 4 = Pores
from surface of capsule; the arrows point up in capsule, X 365. 5 = Cross sections
of a few lamellae showing their height, and depth of notching of apical cell, X 250.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
319
On soil. — Unalaska, Alaska; Lake Lindeman and Dawson, Yukon;
Albert Canon, Selkirk Mountain, British Columbia; eastern British
America; England.
6. Polytrichum jensenii Hag., in Saretryk Middel. om Groenl.
15, p. 444, (1898).
Polytrichum fragilijolium Lindb. fil., in Proc. Soc. pro Fauna et
Fl. Fenn., Oct. 1900.
Fig. 24. Polytrichum jensenii.
1 = Moist plant, XL 2 = Dry plant, XL 3 = Leaf showing lamellae and
margin, X 15. 4 = Leaf tip, X 65. 5 = Cross section of leaf showing lamellae, X
65. 6 = Cross section of a few lamellae, X 250.
Proc. Wash. Acad. Sci., August, 1910.
320
FRYE
Named after Jensen.28
Plants lighter in color than P. commune. Stem 3-5 cm. high.
Leaves about 8 mm. long, somewhat spreading when moist, but
erect when dry, sheath to blade about as i : 2 ; apex blunter than in
P. commune, dentate; a brown spot where sheath joins blade near
vein. Margin plane below, erect or slightly incurved above, en-
tire below, somewhat serrate in the upper half. Lamellae 30-35,
6-8 cells high; marginal cells notched in cross section, smooth,
thickened.
Capsule 4-angled, width to length about as i : ii, smooth, porose.
Teeth 64. Lid conic, with a short slightly crooked beak.29 — In
Sphagnum bogs and on soil. — Nome, Alaska; Yellowstone National
Park, Wyoming; Greenland; northern Europe.
Fig. 25. Polytrichum yukonense.
1 = Plant, X i. 2 and 3 = Leaves, showing lamellae on upper side, X 13. 4 =
Cross section of leaf, X 34. 5 = Cross section of a lamella, showing notched marginal
cell, X 135. 6 = Portion of a lamella as seen from side showing irregularity of cell
form and arrangement, X 270. (After Cardot & ThSriot.)
38 C. O. E. Jensen is a Danish apothecary at Kirke Hvalsd, Denmark.
29 This is not a description of the plant, but a noting of some of the characters from
specimens and verification of points from Asa Gray Bulletin and from Bryologist,
4; p. 26. Literature containing Hagen's description was inaccessible to the writer.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
321
7. Polytrichum yukonense C. & Ther. in Proc. Wash. Acad. Sci.
4: 329 (1902).
Named after its locality, the Yukon river in Alaska.
Plants 5-8 cm. high. Stems simple or nearly so, rather naked
below, base covered with white tomentum.
Leaves rigid, when dry suberect, when damp erecto-patent, 4-6
mm. long, i mm. broad, shortly linear-acuminate from an appressed
yellowish sheath. Margin erect, entire. Lamellae about 30, mar-
gins crenulate, 8-12 cells high. Vein excurrent as an opaque red
entire arista; marginal cells higher than the others, deeply grooved.
Otherwise unknown. — Yukon River, Alaska.
Fig. 26. Polytrichum sexangulare.
1 = Moist plant, X i. 2 = Dry plant, XL 3 = Capsule with calyptra, immature,
X 5. 4 = Capsule, X 5. 5 = Peristome, X 150. 6 = Leaf showing lamellae on upper
surface, X 15- 7 = Leaf tip, showing teeth at back, and involute margins making it
cucullate, X 65. 8 = Cross section of leaf showing lamellae, X 65. 9 = Cross section
of a few lamellae, X 250. (Nos. 6 and 7 after Dixon and Jameson.)
8. Polytrichum sexangulare Floerke, in Hoppe Bot. Taschenb.
1799. p. 126.
Name derived from sex = six, and angularis = angled; referring
to the 6-angled capsule.
Plants 2.5-10 cm. high, in tufts or loose patches. Stems erect or
decumbent, simple or slightly divided, rigid, not tomentose at base.
322 FRYE
Leaves short, rather obtuse, linear-lanceolate from a broad sheath-
ing base, patent when moist, lower ones glossy and dark colored;
dry leaves closely imbricate, rigid, curved at apex. Margin entire,
incurved from near base of limb, at apex cucullate, blunt. Lam-
ellae 30-50, 4-6 cells high; marginal cells larger than the others,
ovate-conical, smooth.
Calyptra not covering the whole of the capsule.
Capsule erect or inclined, oval, with 6 obtuse angles, reddish
brown; length to width about as i : if ; hypophysis obconic, not
constricted above. Teeth short, unequal. Lid with rather thick
beak. Pedicel 1.3-4 cm. long, short, rather thick. — On soil and
rocks. — Rocky Mountain region of British Columbia; Mt. Rainier,
Washington; Mt. Hood, Oregon; Greenland; Europe; Kamchatka
Peninsula; Herald Island, northwest of Bering Strait.
9. Polytrichum juniperinum Willd., in Fl. Berol. Prodr. p. 305.
(1787).
So named because its leaves resemble those of the juniper.
Plants scattered, 2-10 cm. high, glaucous green; male plants more
slender, with short leaves. Stems rarely branched, sometimes
slightly tomentose at base.
Leaves less crowded at top of stem than in P. piliferum, spreading
when moist, erect when dry, long, lanceolate from an oblong base.
Margin incurved but not so nearly meeting as in P. piliferum, en-
tire. Lamellae 35-40, 4-7 cells high; marginal cells longer than
wide, ovate or flask-shaped, smooth. Vein excurrent in a red den-
tate sometimes slightly hyaline arista which is f-if mm. long,
strongly toothed at back of apex, and often half way down. Peri-
chaetial leaves longer than foliage leaves, with longer arista, white
and membraneous at edges.
Calyptra covering capsules.
Capsule larger than in P. piliferum, sharply 4-angled, oblong;
hypophysis short, less distinct than in P. piliferum. Lid deep
red, beak short. Pedicel 4-6 cm. long, shining, bright red. — On
soil. — From Kotzebue Sound north of Bering Sea along the coast
to California; Rocky Mountains; eastern North America; Green-
land. Our most common moss in this family.
THE POLYTRICHACE^E OF WESTERN NORTH AMERICA
323
Fig. 27. Polytrichum juniperinum.
1 = Dry plant with capsule, X i. 2 = Moist antheridial plant; d = antheridial
disk, X i. 3 = Immature capsule with calyptra, X 5- 4 = Capsule, X 5. 5 =
Peristome, X 150. 6 = Leaf showing incurved margin and lamellae on upper side,
X 15. 7 = Leaf tip, X 65. 8 = Cross section of leaf, X 65. 9 = Cross section of
a few lamellae showing smooth flask-shaped marginal cells, X 250.
t
\
10
Fig. 28. Polytrichum strictum.
1 and 2 = Moist plants with capsules, XL 3 = Dry antheridial plant, XL 4
= Dry archegonial plant, XL 5 = Immature capsule with calyptra, X 5- 6 and
7 = Capsules, X 5. 8 = Peristome, X 150. 9 and 10 = Leaves showing lamellae on
upper side, X 15. 11 and 12 = Leaf tips, X 65. 13 = Cross section of leaf, X 65.
14 = Cross section of a few lamellae showing flask-shaped marginal cells, X 250.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA 325
10. Polytrichum strictum Banks, in MSS.
Polytrichum behringianum Kindb.,30 in Rev. Bryol. 1894, p. 39.
Strictum = drawn tightly; probably referring to the closely ap-
pressed leaves in the dry plant.
Plants 6-20 cm. tall, densely tufted, rigid, almost terete when dry
with its closely appressed leaves. Stems more slender than in P.
juniperinum, usually covered for the greater part of its length
with dirty-white tomentum, slightly branched.
Leaves erect-patent, shorter and narrower than in P. juniperinum,
straight, closely and regularly imbricated when dry. Margin in-
curved, entire. Lamellae 25-35, 4~7 ceUs high; marginal cells longer
than wide, ovate or flask-shaped, smooth. Vein excurrent in a red
dentate arista.
Calyptra covering the capsule.
Capsule sharply 4-angled, cubic or very little longer than wide;
hypophysis short. Pedicel 4-6 cm. long. — On soil. — From Kotzebue
Sound north of Bering Strait southward along the coast to British
Columbia; Cascade and Rocky Mountains of British Columbia;
northeastern North America; Greenland; Europe; Asia; Argen-
tine Republic.
11. Polytrichum hyperboreum R. Br., in Parry, Voy. Suppl. p.
294, (1824).
Polytrichum boreale Kindb., in Laubm. Schwed. U. Norw. (1883).
Name derived from hyper = beyond, and boreas = the north
wind; referring to its arctic habitat.
Plants dioicous, 3-12 cm. high. Stems simple or branched,
branches in tufts.
Leaves spreading when moist, appressed when dry. Margin
broad, incurved, entire. Lamellae more or less crenulate, 5-7 cells
high, 25-35; marginal cells ovate or flask-shaped, higher than wide,
smooth, larger than the others. Vein excurrent in a very thick,
short, hyaline, slightly serrate arista.
Capsule erect or inclined, sharply 4-angled, papillose, about if
times as long as wide; hypophysis flattened, deeply constricted from
30 A comparison of authentic Polytrichum behringianum with Polytrichum strictum
shows them to be the same.
326
FRYE
the capsule. Lid hemispheric, with short beak. Pedicel 8-12 cm.
long. — St. Paul Island, Bering Sea; Lake Lindeman, Yukon, Brit-
ish America; Labrador; Greenland; northern Europe and the Alps;
Siberia.
Fig. 29. Polytrichum hyperboreum.
1 = Moist plant, XL 2 = Dry plant, XL 3 = Capsule, X $. 4 = Peristome,
X 150. 5 = Leaf showing lamellae and involute margin, X 15. 6 = Leaf tip, X 65 .
7 = Cross section of leaf, X 65. 8 = Cross section of a few lamellae.
THE POLYTRICHACE.E OF WESTERN NORTH AMERICA
327
Fig. 30. Polytrichum pilifenun.
1 = Moist plant with capsule, XL 2, 3 and 4 = Dry plants, X i. 5, 6 and 7 =
Capsules, X 5. 8 = Capsule with calyptra, immature, X 5. 9 = Peristome, X
150. 10 and 11 = Leaves showing involute margin and lamellae on upper side, X 15.
12 and 13 = Leaf tips, X 65. 14 = Cross section of leaf, X 65. 15 = Cross section
of a few lamellae showing flask-shaped, somewhat thickened cells, X 250.
328 FRYE
12. Polytrichum piliferum Schreb., in Spic. fl. Lips. p. 74, (1771).
Polytrichum pilosum Neck., in Meth, M. p. 123, (1771).
Polytrichum laevipilum Hampe, in Linn. 1859. p, 459.
Polytrichum piliferum var. hoppei Rab.31 in Deutsch. Kryptfl. II, P.
Ill, p. 238 (1848).
Name derived from pilus = hair, and ferre = to bear; referring
to the long hyaline arista at the tip of the leaf.
Plants in loose tufts, glaucous green, 2.5-4 cm. high. Stems
erect, simple, rarely forked, naked at base.
Leaves when dry closely appressed and straight forming a narrow
ovoid or clavate head, leaf-base longer and narrower than in P.
sexangulare, limb narrowly lanceolate, apex below arista minutely
scabrous; leaves of antheridial plants shorter and more shortly
aristate. Margins broadly inflexed and almost meeting, entire,
of very narrow transversely elliptical cells. Lamellae about 30, 4-7
cells high; marginal cell larger than the others, ovate or flask-shaped,
not papillose. Vein reddish, at apex suddenly becoming hyaline,
excurrent as a long denticulate hyaline arista, smooth at back.
Perichaetial leaves longer than the foliage leaves; inner ones thin,
whitish, without lamellae, longly aristate.
Calyptra covering the capsule.
Capsule erect, small, inclined when dry, shortly oblong, with 4
sharp angles and occasionally a fainter intermediate one; hypophy-
sis indistinct, short, constricted above where it joins the capsule.
Lid red or orange, shortly and stoutly beaked. Pedicel 2.5-4 cm.
long. — On rocks and soil. — Lake Lindeman, Yukon, British America;
Vancouver Island and eastward in British Columbia to the Rocky
Mountains; Washington; California; Uinta Mountains, Utah;
eastern British America; Greenland; Europe; Asia; South America.
11 The distinctions between P. piliferum and its variety hoppei do not seem to war-
rant a variety. For example, in Fig. 30, Nos. i, 2, 5, and 12, from the same plant, are
variety hoppei in leaf tips, plant form and capsule; but the nodding capsule does not
agree. Nos. 4, 6, and 13, from another package are variety hoppei in form of capsule
and possibly in leaf arrangement, but hardly in leaf tips; also disagreeing in that many
of the capsules in this material not shown are nodding. No. 7 is not variety hoppei
in form, but the leaves in this plant were very long awned. The length of the awn does
not vary constantly with the form of the capsule nor with the length of the leaf. It is
believed therefore that variety hoppei is P. piliferum prematurely dried or growing
under very adverse conditions, and varying sometimes in capsule, som Jtimes in leaf.