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BFNA Title: Hypnum |
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Hypnaceae, Hypnum Edit level R Brum+ XXX. HYPNUM Hedwig, Spec. Musc. Frond. 236. 1801, conserved name * [Greek hypnos, sleep, alluding to ancient use as filler for cushions] W.
B. Schofield Breidleria
Loeske; Drepanium C. Jens.; Pseudostereodon
(Brotherus) Fleischer; Stereodon
(Bridel) Mitten Plants
small to robust, 0.5-- -15 cm, 1--2-pinnate or
irregularly branched; pseudoparaphyllia filamentous to foliose, toothed or
blunt; axillary hairs 3--4-celled. Stems with or without hyalodermis,
with or without central strand. Leaves
of stem and branch similar but branch leaves tend to be smaller and
narrower and with alar cells less well differentiated; broadly to narrowly
ovate; margins sinuate to entire proximally, toothed to entire distally,
sometimes recurved near base and usually plane distally; apex acuminate or
acute; costa double or obscure, usually confined to proximal 1/4 of leaf;
leaf cells usually smooth, usually elongate and somewhat vermicular, alar
cells often differentiated as a distinct group of enlarged or diminished
cells, often shorter than those of the rest of the leaf, indentation just
above the alar region sometimes present;
axillary hairs with 1-4 (usually 1) short brownish basal cells and 2-5
(usually 3 and sometimes more) elongate hyaline distal cells; in most cases
the walls thin and unornamented. Sexual condition autoicous, dioicous
or phyllodioicous; inner perichaetial leaves erect, ovate to lanceolate or
subulate, abruptly narrowed to a slender acumen, serrate or entire, plicate or
not, outer leaves reflexed, costa single, double, or absent. Seta
smooth, yellowish to reddish. Capsule
erect, inclined or horizontal, varying from long-cylindric to ovoid,
usually curved, annulus 1--3 seriate to scarcely differentiated; operculum
conic to rounded-mammillate; peristome double, exostome teeth
subulate-acuminate, yellowish to brownish, outer surface with distinct zigzag
line and lamellae, finely cross-striolate basally, hyaline and papillose
distally, inner face trabeculate; endostomial segments about as high as
exostomial teeth, pale and yellowish, carinate, weakly to strongly split
between articulations, minutely papillose, cilia 1--3 or sometimes
rudimentary. Calyptra cucullate, naked. Species ca. 50 (22 in the flora): widely distributed in
all continents but SELECTED REFERENCES
Ando, H. 1972. Studies on the Genus Hypnum. I. The genus Hypnum
remains a repository for a number of discordant elements, some considered to
belong to other families. The genus once included a high proportion of the
pleurocarpous mosses. The generic
concept accepted here contains some species probably not Hypnum, but
it is a reasonably “natural” genus that can be recognized in the field,
especially when examined with a hand-lens.
I have included Pseudostereodon,
Drepanium, Breidleria and Stereodon within Hypnum, and remain unconvinced of arguments that they should be
segregated. Gametophytic features are
supreme in separation of species although the presence of sporophytes
provides additional features that strengthen the present species concept. Size, branching patterns, leaf form and arrangement,
areolation, especially alar cell differentiation, leaf margins,
pseudoparaphyllium morphology, and stem anatomy are all features used here
for taxonomic distinctions. With further analysis is possible that axillary
hair morphology will prove to be a useful additional character to separate
species. Unfortunately many of these features are altered by environmental
extremes, especially in moisture and light.
In spite of this plasticity, most specimens can be named with
reasonable confidence; there are, however, some specimens that cannot. The problems in discriminating among the
species have been considerably reduced through the elegant studies of H.
Ando, but he treated only part of the genus in detail. Fortuitously, however, he studied most of
the species that occur in To use the key, a number of recommendations, if followed,
are likely to yield more satisfactory determinations. It must be noted again that some plants may
be impossible to name satisfactorily.
Leafy plants of vegetative specimens may consist almost entirely of
branches, with different but characteristic stem leaves unrepresented; this
can happen when the plants form turf-like colonies or when the stem
disintegrates, leaving only branches. Plants that grow in extreme
environments, especially with respect to moisture, may be unusual; for
instance, plants of deeply shaded or wet sites may be markedly
attenuated. Plants in highly
illuminated and well-drained sites are also subject to morphological
alteration that can “mimic” another taxon. Immature specimens can lack
critical characters or are inconsistent in form. Intermixed material can
contain several taxa, and it is necessary to tease out as complete a plant as
possible for determination. Thoroughly wet leafy shoots and remove branches before
removing many leaves from the main stem and retaining the stripped shoot for
determining form of the pseudoparaphyllia. Observations of leaf morphology
should be based on many leaves. All the sections are on wet material in a
drop of water on a microscope slide on a stage of a dissecting microscope.
Remove leaves from the shoot by stripping them toward the stem base using
fine-tipped forceps as aided by low magnification of a dissecting
microscope. It is important to get
complete leaves, including the cells at the point of attachment to the
stem. In hyalodermous stems, the
thin-walled cortical cells usually remain attached to the leaf base when the
leaves are stripped from the stem; cross-sections can be made of the stem to
confirm whether the stem is hyalodermous or not. Use sporangium-bearing
specimens when possible to provide additional characters. In this key, the
following features define differences in size of gametophores: small refers to stem
leaves 1.2--1.5 mm and stems usually 1--2 cm and slender (related to the leaf
size); medium refers to stem leaves 1.5--3 mm and
stems 2--4 cm; large or robust refers to stem leaves more than 3 mm and stem
more than 4 cm. When length and width of leaves are given, length is cited
first, and the width is that of the widest part of the leaf, usually the base 1. Leaves stripped from stem bearing thin-walled cortical
cells; stems usually with hyalodermis. 2. Hyalodermis
absent, but sometimes an interrupted epidermis of thin-walled cells,
especially at leaf decurrencies .. 8. Hypnum
fauriei (in part) 2. Hyalodermis present as a cortical layer. 3. Stem leaves gradually tapering to insertion, alar
cells only slightly differentiated. 4. Plants somewhat flattened; with leaves not strongly
secund, leaf apex broadly acute. . . ...................................................................................................... 17.
Hypnum pratense 4. Plants never flattened; leaves falcate-secund, leaf
apex narrowly acuminate. 5. Plants
small (stem leaves 1.3--1.5 x 0.4--0.5 mm), usually erect and tuft-forming,
laminal cells 40--60(--70) x 3--4 \um in lumen. . 10..Hypnum hamulosum 5. Plants
medium-sized (stem leaves 1.5--2.8 x 0.4--0.6 mm) usually creeping and not
tuft-forming, laminal cells 50--80(--90) x 3--4 \um; leaf acumen with sharp
teeth . . . 21. Hypnum subimponens 3. Stem leaves curved to insertion, sometimes auriculate,
alar cells usually well differentiated. 6. Plants
small (stem leaves 1--1.5 mm), stem leaves strongly curved to insertion,
sometimes auriculate, alar cells differentiated in a small group at
insertion………16. Hypnum plicatulum 6. Plants small to large, stem leaves weakly to strongly
curved to base; alar cells forming a well-defined group, with the outermost
cells thin-walled. 7. Plants small to medium-sized; stem leaves broadly
ovate lanceolate, alar cells gradually enlarged downward, never decurrent . .
. 11. Hypnum holmenii 7. Plants
medium-sized to large; stem leaves ovate to oblong lanceolate; alar cells
abruptly differentiated into well-defined group that is thin-walled, swollen
and sometimes decurrent. 8. Alar
cells weakly decurrent, usually not pronounced on dissection, 2--3
rectangular cells vertically and 3--4 cells horizontally, the outermost basal
cells thin-walled and often collapsed inward, with a few triangular cells
above the outer hyaline cell and cells beyond the thin-walled cells with
thickened, often pigmented walls; stem red-brown to nearly black.. . . 7. Hypnum dieckii 8. Alar
cells in a conspicuous decurrent group of thin walled, often swollen cells
stripping off on dissection, without thicker-walled pigmented cells; stems
yellow or reddish. 9. Plants medium sized to small, usually reclining,
leaves with long-filiform serrulate acumen, stems yellow or green… . . 3. Hypnum callichroum 9. Plants
large, often suberect, leaves with acute or broadly acute apex, older stems
reddish . . 14. Hypnum lindbergii 1. Leaves stripped from stem without thin-walled cortical
cells; stems lacking hyalodermis, but occasionally with scattered thin-walled
cortical cells. 10. Leaves with margin strongly revolute from the base to
upper portion . . . 20. Hypnum revolutum
(in part) 10. Leaves with margin plane or sometimes recurved near
base. 11. Stem leaves auriculate at insertion, auriculation
sometimes only on one side. 12. Stem
leaves with long-attenuate apex; cells below auricle enlarged and frequently
reddish pigmented, much enlarged, alar cells few (5--6) in auriculation . . .
4. Hypnum circinale 12. Stem
leaves gradually tapering to a point, not long-attenuate; cells below
auriculate portion not conspicuously enlarged, or absent; alar cells numerous
(more than 20) or reduced to less than 10. 13. Alar
cells more than 20, not excavate; plants brownish, not glossy; plants closely
pinnately branched . . . 18. Hypnum procerrimum 13. Alar
cells less than 10, often excavate; plants yellowish to golden green, glossy;
plants distantly pinnately branched . . . . . 6. Hypnum curvifolium 11. Stem leaves gradually curving to insertion, never
auriculate. 14.
Many stem leaves when stripped from stem bearing one or more elongate
decurrent cells on margin. 15. Pseudoparaphyllia foliose or lanceolate, with incised
margins… 12. Hypnum imponens (in
part) 15. Pseudoparaphyllia foliorse or triangular to
lanceolate, entire . . . 8. Hypnum fauriei (in part) 14. Stem leaves usually stripping from stem not bearing
elongate decurrent cells. 16. Alar cells forming an extensive triangular area of 5
or more vertical rows of quadrate cells and few or no elongate cells at
insertion. 17. Pseudoparaphyllia broadly foliose, blunt. . . 22. Hypnum vaucheri (in part) 17. Pseudoparaphyllia filamentous, lanceolate or lobed,
sharply pointed. 18. Stem
leaves strongly recurved near base and up margin; autoicous, sporophytes
frequent . . . 19. Hypnum recurvatum 18. Stem leaves weakly recurved or plane near base;
dioicous 19. Plants
slender, branch leaves (0.8--)1--1.4 x 0.2--0.4 mm. 20. Stem
leaves falcate-secund, subentire to strongly toothed in long-attenuate apex .
. . .............. 1. Hypnum andoi (in part) 20. Stem
leaves weakly or not falcate, entire to weakly serrulate in apex . . . 5. Hypnum
cupressiforme (in part) 19. Plants
medium to large in size, branch leaves larger. 20. Plants
robust, branch leaves 1.4--2.5 x 0.5--0.7 mm 20. Plants
medium-sized, branch leaves 1.4--2 x 0.4--0.5 mm. 21. Plants
pale green, strongly complanate, leaves loosely imbricate . . . 13. Hypnum jutlandicum (in part) 21. Plants
greenish to golden or rusty green, never complanate, leaves strongly
imbricate . 5. Hypnum cupressiforme
(in part) 16. Alar
cells forming a small region of 1--3 rows of quadrate cells with 1-3 elongate
cells at the insertion. 22. Pseudoparaphyllia filamentous to lanceolate. 23. Plants
pale green, strongly complanate, leaves loosely imbricate . . . 13. Hypnum jutlandicum (in part) 23. Plants
rusty to dark green to yellowish, never complanate, leaves closely imbricate. 24. Stem
leaves falcate-secund, leaf apices strongly toothed, plants usually hanging
downward on perpendicular substratum . . . 1. Hypnum andoi (in part) 24. Stem
leaves falcate secund or straight, leaf apices entire or nearly so, plants
usually creeping over horizontal substratum . . . 5. Hypnum cupressiforme (in part) 22. Pseudoparaphyllia foliose. 25.. Alar
cells usually in dark brown group, often with an excavate group above the
insertion; cells pitted throughout; plants usually dark chestnut brown and in
seepage sites . . . 2. Hypnum bambergeri 25. Alar
cells, if pigmented, usually reddish to yellowish, if excavate, the whole
alar region involved; cells pitted only near base; plants pale rusty brown to
golden green, usually in well-drained to dry sites. 26. Plants slender, alar cells unpigmented. 27. . Stem
leaves toothed near apex, autoicous, often with sporophytes, epiphytic or xilicolous.
. . 15. Hypnum pallescens 27. Stem
leaves entire near apex, dioicous, sporophytes absent, terrestrial and calcicolous
. . . 20. Hypnum revolutum (in
part) 26. Plants
medium-sized to robust; dioicous, alar cells pigmented or not. 28. Alar
cells not pigmented or swollen at insertion. . . 22. Hypnum vaucheri (in part) 28. Alar
cells with red-pigmented somewhat elongate swollen cells in 1--4 rows at
insertion. 29. Pseudoparaphyllia
bearing several sharp 1-seriate teeth …
12. Hypnum imponens (in part) 29 . Pseudoparaphyllia
lanceolate, or if lobed, lacking additional teeth. . . 9. Hypnum
fujiyamae 1.
Hypnum andoi A. J. E. Smith, J. Bryol 11: 606. 1981 (1982) Hypnum cupressiforme
var. mammillatum Bridel; H. mammillatum (Bridel) Loeske,
illegitimate name Plants slender,
pale green to yellowish green, 2--8 cm, reclining. Stems small, pale yellowish brown, creeping, closely affixed to
substratum, epidermal cells not hyalodermous, central strand weakly
developed; usually irregularly to regularly pinnately branched, branching
arising in horizontal plane, pseudoparaphyllia subfilamentous to lanceolate,
tipped by elongate cell. Leaves weakly to strongly
falcate-secund, ovate- or oblong-lanceolate, tapering gradually to base and
tapering gradually to a short or more usually long acumen, 1--1.8 x
0.35--0.60 mm, margins usually sharply serrulate, often recurved near base,
costa double, short or indistinct; median cells (40)50--60(--70) x 4--5 \um,
basal cells shorter and wider, usually yellow or yellowish brown, alar region
weakly excavate, often brown, of subquadrate cells (5--)7--10(--13) along
margin, the marginal ones enlarged and hyaline, the inner brownish. Sexual condition dioicous; inner perichaetial leaves erect,
oblong-lanceolate, suddenly narrowed to a long, serrulate acumen, costa obscure. Seta
yellowish to reddish brown 0.08--1.7(--2) cm. Capsule yellowish to reddish brown, erect to inclined,
oblong-cylindric 1.5--1.8(--2) mm; operculum conic (rounded-mammillate);
annulus 2--3 seriate; cilia of endostome single or imperfectly double, very
fragile. Sporophytes
produced in late autumn (September), capsules often maturing in spring
(Mar.--Apr.), infrequent.
Perpendicular surfaces of tree trunks and rock cliffs; 0--2000 m; Hypnum
andoi is a temperate species of amphi-Atlantic distribution.
This species resembles Hypnum
cupressiforme var filiforme,
but the leaves of H. andoi are long attenuate, and the row of basal
hyaline cells in the alar region is decidedly different, reminiscent of Sematophyllum; the shorter, stouter
capsule, and mammillate operculum are also valuable characters but
sporophytes are extremely rare in North America, and have a conic operculum
rather than a mammillate one, as in Europe. 2.
Hypnum bambergeri
Schimper, Syn. Musc. Europ. (ed.2) 698. 1860 Stereodon bambergeri (Schimper)
Lindberg Plants robust,
2--8(--10) cm, ascending to prostrate, reddish to yellowish brown, shiny,
usually little and irregularly branched, but occasionally pinnately branched,
with branches of variable length 0.2--0.6 cm, leafy stems 0.1--0.2 cm broad. Stems dark brown to reddish brown,
with weak central strand; pseudoparaphylla wide, foliose. Leaves of stem leaves falcate to
circinate-secund, oblong-ovate to oblong-lanceolate, narrowed to a slender
acumen, margins plane and sinuate to weakly serrate, 1.5--2 x 0.4--0.6 mm,
curving gradually to alar cells, or forming a weak auricle just above the
alar cells; costa single or double, and unequal, short to long, unequal;
median cells 30--60 x 35--6 \um, somewhat to markedly pitted, cells of leaf
base shorter and more strongly pitted, pigmented yellow to orange; alar cells
in a well-defined, weakly to markedly excavate brownish group 3--7 cells high
along margin and extending to 3--6 cells wide, quadrate to
short-rectangular. Sexual condition
dioicous; inner
perichaetial leaves erect, ovate to oblong-lanceolate, abruptly narrowed to a
slender entire acumen, laciniate at shoulders, plicate. Seta
reddish brown, 1.3--2 cm. Capsule yellowish brown, inclined to
horizontal, 1.3--1.5 mm, excluding the conic operculum, cilia of endostome
2--3 Sporophytes
infrequent, capsules mature summer (July, August). Often in seepage habitats,
also in dry tundra, open conifer forest, mainly calcicolous; 0--2500 m;
Greenland; Alta., B.C., Man., Nfld., N.W.T., Nunavut, Ont., Que., Yukon;
Alaska, Mont.; Europe; Asia. Hypnum
bambergeri is widely distributed in the 3.
Hypnum callichroum
Bridel, Bryol. Univ. 2: 631. 1827 Hypnum
alaskae Kindberg; Stereodon
callichrous (Bridel) Braithwaite . Plants medium-sized,
2--6 cm, branches 0.2--1 cm, regularly to irregularly pinnate, pale green to
yellow-green, shiny, procumbent to ascending, firmly or loosely attached to
substratum. Stems yellowish to green, hyalodermous, with central strand,
pseudoparaphyllia lanceolate. Leaves of stem falcate to circinate secund
ovate to ovate-lanceolate gradually narrowing to a slender acumen, 1.5--2 x
0.75--0.8 mm, somewhat curved distal to alar region, margins plane, often sharply
toothed in acumen that tapers to a 1-seriate apex, costa short and double or
obscure, median cells linear, 50--60 x 8 \um, basal cells shorter, broader
and porose, yellowish to unpigmented; alar cells forming a well defined
region of thin-walled, often somewhat inflated cells 1--3 cells high and
bounded by quadrate to triangular cells above. Sexual condition dioicous; inner perichaetial leaves erect,
oblong-lanceolate and finely pointed, plicate, toothed near apex, costa
obscure. Seta 2--3.5 cm, red-brown, capsules red-brown, inclined, somewhat
curved, 1.5--2 mm excluding conic operculum, annulus 1--2 seriate. Sporophytes
produced summer, capsules maturing June--July. Epiphytic or more usually
terrestrial, forest, open terrain; 0--1500 m; Greenland; Alta., B.C.; Alaska,
Wash.; Europe; Asia. In
4.
Hypnum circinale Hooker, Musci Exot. 2: 21. 1820
Hypnum
squoitei J. K. A. Müller; Rhaphidostegium recurvans
Kindberg; Stereodon circinalis (Hooker)
Mitten Plants light
gray- to golden-green or dark green, 3--5(--10 or more) cm. Stems slender, irregularly to
regularly pinnate (occasionally 2-pinnate) or irregularly branched, usually
creeping and firmly affixed by rhizoids, reddish brown, lacking hyalodermis
or central strand; pseudoparaphyllia filamentous to lanceolate, usually
terminated by an elongate cell or 1-seriate tip 2--4 cells long. Leaves of stem falcate-secund to circinate, ovate- to triangular-lanceolate,
often asymmetric at base with one side, somewhat to strongly auriculate,
slightly decurrent, narrowing to a long acumination, 1.5--2.2 x 0.5--0.7 mm,
margins serrulate, plane, rarely slightly recurved on one side, costae
indistinct; branch leaves smaller, 1.1--1.5 x 0.3--0.4 mm, more strongly
serrulate, median cells 60--80(--100) x 4--5 \um, with thin or thick porose
walls, basal cells broader with porose walls, golden yellow, alar cells few,
subquadrate to rounded-triangular, 2--5 in marginal row, proximal ones
inflated, reddish brown, sometimes hyaline on margin. Sexual condition dioicous;
antheridial plants similar to archegonial or phyllodioicous with dwarf males epiphytic on archegonial
plants; perichaetial leaves erect,
oblong-lanceolate, inner erect, the outer recurved, slender point serrulate,
not plicate. Seta smooth, reddish brown, 0.6--1.5(--2) cm. Capsule reddish brown, oblique to
horizontal, ovoid to ovoid-oblong, 0.8--1.5 x 0.5--0.7 mm, operculum
conic-apiculate, annulus 1--2 seriate; endostomial cilia 1--2. Sporophytes
produced autumn--winter (Sept.--Dec.), capsules usually mature
Jan.--Feb. Lowland to subalpine
coniferous forests, commonly epiphytic on tree trunks, also on decaying logs
and rock; 0--1500 m.; endemic to western North America; Alta., B.C.; se
Alaska, Calif., Idaho, Mont., Oreg., Wash. The
often asymmetric stem leaves bearing a long-attenuate serrulate point, with
one side auriculate with usually colored alar cells, the small sporangia that
mature in early spring make Hypnum circinale an easy species to
recognize. Its closest affinities are
with the east Asian H. tristoviride
(Brotherus) 5. Hypnum cupressiforme Hedwig,
Spec. Musc. Frond. 291. 1901 Hypnum
pseudo-fastigiatum J. K. A. Müller; Stereodon cupressiformis (Hedwig)
Mitten Plants rusty
green, golden green, yellow-green to pale green, medium sized to small, 2--10
cm, occasionally longer, prostrate to suberect. Stems pale to yellowish green, brown with age, epidermal cells
not hyalodermous, central strand poorly developed, irregularly pinnate to
nearly unbranched, shoots, attached shoots often regularly pinnate; pseudoparaphyllia
filamentous, 1--3-seriate at base, but ending in elongated pointed 1-seriate
tip. Leaves weakly to decidedly falcate-secund, ovate to
oblong-lanceolate, generally gradually curved to insertion and tapered to a
slender acumen, 1.5--2 x 0.5--0.8 mm, margins serrulate (sometimes weakly) in
distal half, occasionally nearly entire, recurved to plane proximally; costa
double and short to obscure; median cells (50--)60--80 x 3--4(5) \um; basal
cells shorter and wider; alar regions weakly or not excavate, made up of a
well-defined triangular to quadrate area of many subquadrate cells or reduced
to very few quadrate cells (sometimes pigmented), the more basal cells
larger, sometimes hyaline, yellowish or brownish. Sexual condition dioicous; inner
perichaetial leaves erect, oblong-lanceolate, serrulate distally, not
plicate; margins plane; costa obscure. Seta reddish brown, 1--2.5(--3) cm. Capsule reddish brown, slightly
inclined, cylindric and slightly curved, 1.8--2.5(--2.8) mm, excluding conic
to rostrate operculum, annulus
1--3-seriate; cilia of endostome usually 1--2(--3). Varieties
9 in the world (4 in the flora):
Cosmopolitan; absent in Hypnum cupressiforme is
an extremely polymorphic species, reflected in the more than 60 varieties that
have been described. It has a wide ecological amplitude as well as a
cosmopolitan world distribution and is found in all climatic regions.
Taxonomic features reliable in most other species of Hypnum are plastic in H.
cupressiforme. Within a single clone it is possible to sort out several
named varieties. Hypnum cupressiforme
var. lacunosum Bridel was noted by
H. Ando (1989) to be weakly differentiated in North America, and tentatively
cited specimens from 1. Plants small, branch leaves 1--1.4 x 0.2-0.4 mm or
slightly less. 2. Leaves straight to slightly
falcate, entire to weakly toothed distally. 2. Leaves falcate-secund, subentire
to distinctly toothed distally . . 5a. Hypnum
cupressiforme var. cupressiforme (in
part) 3.
Plants subjulaceous to filiform, prostrate, leaves oblong-lanceolate,
gradually narrowed to acumen; cells 60--80 x 3--4 \um (or slightly
greater)……………………. 5b. Hypnum cupressiforme var. filiforme 3.
Plants julaceous, usually in compact tufts; leaves ovate or widely
oblong-lanceolate, abruptly narrowed to short acumen; median cells shorter
and broader, 40--50 x 5--6 \um …. 5c. Hypnum cupressiforme var. julaceum 1. Plants medium-sized to robust, branch-leaves 1.5--2 x
0.4-0.6 mm or slightly greater. 4.
Plants robust, branch leaves 1.4--2 x 0.4--0.7 \um or greater, branches not
or rarely flagellate at tips . . . . [Hypnum
cupressiforme var. lacunosum] 4.
Plants medium-sized, branch-leaves 1.5--2.5 x 0.5--0.8 mm or greater. 5.
Plants weakly complanate, leaves falcate-secund; alar cells not excavate or
colored…5a. Hypnum cupressiforme var.
cupressiforme (in part) 5.
Plants subjulaceous to complanate; leaves straight to weakly falcate; alar
cells often excavate and colored . . . . 5d. Hypnum cupressiforme var. subjulaceum 5a.
Hypnum cupressiforme Hedwig var. cupressiforme Plants
varying from slender to medium-sized and in color from pale green, dark green
to golden-brown; length various; branching pinnate to irregular, reclining to suberect. Leaves usually falcate, sometimes
strongly; alar cells numerous to few, usually not excavate. Producing
sporophytes late summer--fall, capsules often mature spring. Terrestrial,
epiphytic, epilithic; 0--4000 m; Greenland; Alta., B.C., Man., N.B., Nfld.,
N.W.T., N.S., Nunavut, Ont., P.E.I., Que., Sask., Yukon; Alaska, Ark., Colo.,
Conn., Del., Ill., Iowa, Maine, Md., Mass., Mich., Mont., N.Y., N.C., Pa.,
S.Dak., Vt., Tex., Wash., Wis.; Mexico; South America; Eurasia; Africa;
Atlantic Islands; Pacific Islands (New Zealand); Australia. Hypnum cupressiforme var.
cupressiforme is used as a
catch-all to accommodate specimens that cannot be placed in another of the 9
varieties recognized by H. Ando including those accepted here. 5b. Hypnum cupressiforme var. filiforme
Bridel, Musc. Rec 2: 138. 1801 Plants slender,
pale green to dull green, occasionally yellowish green, 2--6 cm, irregularly
to somewhat pinnately branched, some branches frequently filiform. Leaves
subjulaceous, straight to somewhat falcate, gradually narrowed to slender
acumen, nearly entire, alar cells subquadrate, usually in a well-defined
group. Sporophytes unknown in Mainly
confined to perpendicular surfaces of cliffs and tree-trunks; 0--2500 m;
N.B., N.S., P.E.I.; Ark., Conn., Maine, Mass., Mich., N.H., N.J., N.Y., N.C.,
Okla., Pa., Tenn., Tex., Va; South America (Chile); Europe; Asia; Africa;
Pacific Islands (New Zealand); Australia. When
well developed, Hypnum cupressiforme var. filiforme is
distinctive, with filiform julaceous stems and straight leaves. There are variants with somewhat
falcate-secund leaves that can be confused with H. andoi. The latter
species rather has long-attenuate leaf apices, is not julaceous, although
filiform, and the leaves are not strongly imbricate as in var. filiforme. The many specimens that closely resemble
var. filiforme strongly suggest that it may be an environmental form
induced by the habitat that holds moisture rather briefly during the growing
season. 5c. Hypnum cupressiforme var.
julaceum Bridel, Musc. Recent.
Suppl. 2: 216. 1812. Plants tufted,
medium sized, prostate to erect, to 5 cm, densely to loosely pinnately
branched, sometimes asymmetric, branches 0.2--1.5 cm, julaceous. Leaves straight to weakly falcate, ovate
or widely oblong, concave, acumen abruptly narrowed to short-acuminate, subentire. Seta 1--1.5 cm. Capsules ca. 1 mm. Confined
to horizontal, rather than vertical surfaces, strongly illuminated sites,
especially in mountains and at higher latitudes; 100--2500 m.; N.W.T., 5d. Hypnum cupressiforme var.
subjulacuum Molendo, Ber.
Naturhist. Ver. Augsburg 18: 183. 1865 Plants creeping,
medium-sized, yellowish green to brown, lustrous, usually prostrate, 5--8 cm
or more, regularly to irregularly pinnate, to 2-pinnate, occasionally plumose
on one side, branches 0.2--1(--1.5) cm, subjulaceous, sharply pointed. Leaves
of stem oblong-lanceolate,
straight to weakly falcate, sometimes homomallous, gradually narrowed to long
and slender acumen, somewhat concave, margins subentire to weakly toothed,
weakly recurved near base or plane throughout; alar cells subquadrate, often
brownish, often in excavate group. Sporophytes
unknown in area of flora. Terrestrial,
cliff shelves, horizontal rock surfaces, in both exposed and sheltered sites,
predominantly calcareous substrata; 0--4000 m; B.C., Yukon; Alaska, Colo.,
N.Mex., N.Dak.; Europe (mainly high elevations); Asia; Pacific Islands (New
Zealand). This
variety is frequent in the 6.
Hypnum curvifolium Hedwig, Spec. Musc. Frond.
285. 1801 Stereodon curvifolius (Hedwig)
Mitten Plants medium-sized
to robust, procumbent to suberect, light green to yellow-green, generally
regularly pinnately branched, 4--10 cm; branches to 1.5 cm, leafy stems
0.5--3 mm wide. Stems orange brown to reddish, hyalodermis absent,
central strand weakly differentiated; pseudoparaphyllia foliose. Leaves of stem ovate, falcate-secund, 1.5--2.5 x 0.7--0.8 mm, narrowing
to a relatively broad acumen, curving to the insertion, slightly decurrent,
margins plane, sinuose to entire near base, gradually weakly toothed near
apex; costa short and double; median leaf cells 70--80 x 4--5 \um, stouter at
apex and at insertion where broader and pigmented, median cells sometimes
projecting abaxially at distal end of cell, especially near apex; alar cells
few, hyaline and enlarged, in 1--3 rows across base, delimited by smaller
ovoid to triangular cells distally. Sexual condition dioicous; inner
perichaetial leaves erect, lanceolate to subulate, tapering gradually to
apex, somewhat plicate. Seta 2--4.5 cm, orange-yellow to
brownish when mature. Capsule reddish
brown, obovoid-cylindric, 2.5--3 mm excluding conic operculum, strongly
curved, plicate when dry, annulus broad, 2-seriate, cilia of endostome 2--3,
nodulose. Sporophytes
produced in summer; capsules mature July--Aug. Terrestrial or most commonly
on decaying logs in woodland, also rock, occasionally in peatland; 0--2000 m;
N.B., Nfld., N.S., Ont., P.E.I., Que.; Ala., Ark., Conn., Del., D.C., Ga.,
Ill., Ind., Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss.,
Mo., Nebr., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., Tenn.,
Tex., Vt., Va., W.Va., Wis. Hypnum
curvifolium is an eastern North American endemic. It might be
confused with H. lindbergii from
which it differs in the more extensive and often decurrent alar region of H.
lindbergii and the more plumose
branching in H. curvifolium. In Hypnum lindbergii, the leaves are
never as circinate as those of Hypnum
curvifolium. The presence of an
hyalodermis in H. lindbergii and its absence in H. curvifolium
further separates these species. From H. imponens, usually smaller,
the characteristic toothed pseudoparaphyllia of that species separate it. 7.
Hypnum dieckii
Renauld & Cardot, Bot. Centralbl. 44: 423. 1890 Stereodon dieckii
(Renauld & Cardot) Brotherus Plants medium-sized,
4--8 cm, generally regularly pinnate, dark green to golden green or nearly
black, procumbent to suberect, often firmly attached by rhizoids, but when
suberect generally without rhizoids; branches 0.2--1.5 cm, leafy stems
0.05--0.15 cm wide Stems red-brown
to nearly black, hyalodermous, central strand poorly differentiated;
pseudoparaphylla foliose, broad. Leaves
of stem falcate, leaves
curved downward towards substratum, ovate-lanceolate, tapering to a narrow
acumen, 1--1.5 x 0.4--0.5 mm, curving slightly to insertion, margins plane,
entire to sinuate in basal half, but often sharply serrate toward apex, costa
usually indistinct; median cells 40--60 x 3--4 \um, basal cells often
pigmented yellowish to brownish, especially adjacent to hyaline alar cells,
porose, rectangular, shorter and wider than median cells; alar cells in a
well-defined excavate group of thin walled, mainly rectangular cells, area
2--3 cells high and 3--4 cells deep, the outer ones thin-walled and somewhat
collapsed inward, with a few triangular cells distally, often bulging 40--65 x
20--30 \um. Sexual condition dioicous;
inner perichaetial leaves erect, lanceolate and finely pointed, plicate,
toothed, costa obscure. Seta 2--4.5 cm, red-brown when mature. Capsule
red-brown,usually nodding when mature, 3--4.5 mm excluding conic
operculum, annulus 1--2-seriate, cilia
of endostome rudimentary. Sporophytes
produced summer; capsules mature June-July. Terrestrial, rocks, logs, near
water courses, seepage areas, open to somewhat shaded areas, but usually
where persistent moisture is available; 0--1500 m; B.C.; Hypnum
dieckii is distributed around the north Pacific, and can be
frequent on logs or outcrops near streams. This is a distinctive species,
especially when sporophytes are present, as it is the only North American
species that has nodding capsules. The
dark stems, and pinnate branching associated with the excavate region of
thin-walled alar cells set off by inner thick-walled and pigmented basal
cells, are usually enough to separate it.
It can be distinguished from Hypnum
lindbergii by the
decurrent thin-walled alar cells and the longer and weakly toothed leaves of
that species. 8.
Hypnum fauriei
Cardot, Beih. Bot. Centralbl. 17: 41.
1904 Hypnum
fertile of American authors, not Sendt. Plants creeping,
medium-sized, 4--8 cm, ca. 0.13 cm broad including leaves, yellowish brown to
brownish green, somewhat glossy, often densely branched, regularly to
irregularly pinnate, often attached to substratum by numerous rhizoids;
branches up 1 cm, ca. 0.08 cm wide (with leaves). Stems
lacking hyalodermis, reddish to yellowish; pseudoparaphyllia foliose,
triangular to lanceolate. Leaves of stem falcate to circinate,
from broadly ovate base, tapering to a slender acumen and narrowing to
insertion, sometimes weakly cordate, 1.4(--2.2) x 0.4(--0.75) mm, yellowish
brown at base, margin crenate to entire or toothed in acumen, proximal margin
excavate in alar portions where the cells are often colored, hyaline and few
(1--3), rectangular to quadrate and often bulging with 2 or 3 subquadrate to
rectangular cells distally, sometimes with rectangular decurrent cells at
insertion; median laminal cells linear to slightly vermicular, 50--70(--80) x
3--5 \um, basal cells shorter and thicker with thicker porose walls. Sexual
condition autoicous. Seta 3--4 cm, reddish brown or
yellowish brown. Capsule inclined to horizontal, cylindric, 2--3 x 0.9--1 mm,
curved, brown when mature, operculum conic-apiculate, annulus 3-seriate,
cilia of endostome 2--3, endostome segments with large perforations. Capsules
mature summer (July). Tree bases,
stumps, rotten logs, occasionally on humus and rock, mixed forest; 0--2000 m,
lower elevations in the northern part of the range but at higher elevations
in the mountains southwards; N.B., N.S., Ont., Que.; Ga., Maine, Mich.,
Minn., N.H., N.Y., N.C., Pa., Tenn., Vt., Va., W.Va.; E Asia: Korea, Japan,
Russia (Altai). Hypnum
fauriei has posed difficulties since its early report from North
America as H. fertile, a species restricted to 9. Hypnum fujiyamae (Brotherus)
Paris, Ind. Bryol., suppl. 202. 1900 Stereodon
fujiyamae Brotherus, Hedwigia 38: 232. 1899 Plants
robust, prostrate to suberect, 3--15 cm, branches 2--5
cm, sometimes longer, leafy stems 0.2--0.25 cm broad; golden to rusty
green, shiny, irregularly to distantly pinnately branched, stem apices
markedly pointed with imbricate leaves. Stems
yellowish brown, central strand weakly developed; pseudoparaphyllia
lanceolate to ovate, foliose, lacking cilia. Leaves of stem usually falcate-secund, ovate-lanceolate to ovate,
gradually acuminate 0.2--3 x 0.1--1 mm, plicate, acuminate, somewhat toothed,
margins sometimes recurved in proximal half, rarely nearly to apex; costa
double, faint to distinct, basal cells thicker walled and porose,
sometimes brownish yellow, alar cells well defined, forming a group 3--4
cells deep and 5--8 cells wide at base, the inner ones usually colored,
grading into the basal cells, outer cells inflated somewhat and thin-walled,
hyaline to yellowish, grading to brownish, subquadrate to rectangular
supra-alar cells, the leaf margin curved above them, sometimes slightly
auriculate. Sexual condition dioicous or phyllodioicous. Sporophytes lacking in area of flora. Usually tundra fen slopes over shallow peat in late-snow
lying sites; mainly near 200 m; Hypnum fujiyamae somewhat resembles H. lindbergii in the field but the
golden to brownish color of the plants and the imbricate cuspidate apices of
the shoots mark it as different. While H.
lindbergii forms turf-like mats, H.
fujiyamae has reclining shoots. Microscopically these species are clearly
unrelated. 10.
Hypnum hamulosum
Schimper in P. Bruch, W. P. Schimper & W. Gümbel, Bryol. Eur. 6:
96. pl. 590. (fasc. 57--61. Monogr. 20. plate 10: hamulosum). 1854 Stereodon hamulosus
(Schimper) Lindberg Plants small,
yellowish, pinnate to irregularly branched, forming turfs of erect to
suberect shoots or creeping, 2--9 cm, branches 0.4--0.8 cm, slender. Stems
yellowish, with hyalodermis and weak central strand; pseudoparaphyllia
scarce, foliose. Leaves of stem strongly hamate and
falcate-secund, ovate-lanceolate, slightly tapered to base, 1 x 1.5 mm,
margins plane, mostly entire but weakly toothed toward apex; cells narrowly
vermiculate, with blunt ends, 6--7 x 60--75 \um shorter and broader toward
base; alar cells rarely differentiated except at margin and near insertion,
with 2--3 slightly larger, thin-walled cells. Sexual condition phyllodioicous;
perichaetial leaves erect, except the outermost reflexed, plicate, with
attenuate toothed apex. Seta 1--2
cm, reddish. Capsule inclined to
horizontal, long-cylindric, urn 1.5--1.8 mm, operculum conic, 0.5 mm; annulus
2--3-seriate cilia of endostome 1--2, smooth. Capsules
mature July--August, infrequent. Predominantly on calcareous rocks, in
crevices, also reported in tundra, mountains in southern part of its range;
elevation not determined; Alta., B.C., N.W.T., Nunavut, Yukon; Greenland; Alaska,
Colo.(?); Europe; Asia. Hypnum
hamulosum resembles the related species H. callichroum, H. holmenii,
and H. plicatulum, especially the last two, but differs in its strongly
hamate stem leaves that generally do not taper to the base and usually lack
differentiated alar cells. Branch
leaves are not diagnostic, and tundra forms can be impossible to name with
confidence. These often possess only
branch leaves as the main stem is not present. Many reports of H. hamulosum
from 11.
Hypnum holmenii Ando, Hikobia 11. 365. 1994 Stereodon holmenii
(Ando) Ignatov & Ignatova Plants small,
regularly to irregularly and densely pinnately branched, pale green to
yellow-green, shiny, ascending to procumbent, 4--6 cm, branches to 1 cm,
loosely affixed to substratum, turf or weft-forming. Stems
yellowish, hyalodermous, with weak central strand, pseudoparaphyllia
foliose, lanceolate, irregularly dentate.
Leaves of stem falcate, secund broadly
ovate-lanceolate, gradually narrowed to a slender acumen, curved to insertion
(sometimes markedly), 1.4--1.6 x 0.5--0.6 mm, margins plane and serrulate
distally, costa short and double; median cells linear (50--)60--80 x 3--4 \um,
basal cells shorter and wider, yellowish or unpigmented; alar regions are
distinct areas of subquadrate to rectangular cells, 9--20 \um wide, usually
not decurrent, the most basal cells thinner walled and hyaline, often tearing
away with the enlarged stem cortical cells; branch leaves similar. Sexual condition dioicous or
phyllodioicous; perichaetial leaves erect, oblong-lanceolate, with slender
serrulate acumen, plicate, costa obscure.
Seta 1.5--2.2 cm, pale
reddish brown. Capsule red-brown,
inclined to horizontal, oblong-cylindric, curved, 1.5--1.8 mm, excluding
conic operculum; annulus 1--2-seriate, cilia of endostome 2--3. Sporophytes
produced summer; capsules mature July--Aug. Mainly in moist heaths, but also
shrub thickets and spruce forest, largely on calcareous substrata; 0--1500 m
or more; Greenland; Alta., B.C., N.W.T., Nunavut, Yukon; Europe (Finland). Hypnum
holmenii occurs mainly at elevations above 1500 m. In size and
general aspect this species often resembles H. plicatulum, but the leaves are generally not markedly
auriculate, as in H. plicatulum.
The enlarged alar cells are not differentiated in the latter species, while
they usually are in H. holmenii. From Hypnum
callichroum, it differs in the abruptly differentiated group of enlarged
alar cells of the former, these gradually differentiated in H. holmenii. It is necessary to examine many leaves to
achieve an accurate assessment. Hypnum
hamulosum differs in its strongly hamate stem leaves and the less
differentiated alar cells. Because H.
holmenii often resembles related species, its distribution has been
inadequately documented. 12.
Hypnum imponens
Hedwig, Spec. Musc. Frond. 290.
1801 Stereodon imponens (Hedwig)
Mitten Plants golden to yellow-green or brownish, medium-sized to large, creeping, 3--10 cm. Stems reddish brown, central strand weak, hyalodermis not differentiated from the cortical, usually regularly pinnately branched, occasionally partly 2-pinnate, or irregularly branched; branches usually in a single horizontal plane, 0.3--1.2 cm; pseudoparaphyllia frequent, lanceolate or foliose, mostly with inc |