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BFNA Title: Fontinalaceae-Fontinalis |
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XX. FONTINALACEAE
Schimp., Coroll. Bryol. Eur. 96. 1856 Plants slender, medium-sized, or robust,
forming loose or dense masses. Primary
stems creeping. Secondary stems
prostrate, pendent, or trailing, freely and irregularly branched; central strand
absent; rhizoids on stems and at
base of secondary stems, from initials abaxially to the leaf insertions and
around secondary stem primordia; axillary hairs elongate. Leaves 3-ranked, keeled and
conduplicate, concave, or plane; margins entire or serrulate; median leaf
cells linear to linear-rhomboidal; alar cells firm-walled or enlarged and
bulging; costa subpercurrent, percurrent, excurrent to long-excurrent or
nearly absent. Sexual condition dioicous;
perichaetia lateral on short branches. Seta
short or elongate. Capsule
immersed, laterally emergent, emergent or exserted, oval, oval-oblong,
subcylindric or cylindric; stomata absent; operculum obtuse-conic, acute to
obliquely rostrate; annulus rudimentary; peristome diplolepidous, exostome
teeth 16, linear, sometimes united in pairs at tips, outer surface smooth and
lightly thickened, inner surfaces heavily thickened, granulose, at times with
obscure vertical striae, inner trabeculae heavily thickened, endostome
segments linear, red to brownish-orange, papillose, joined by lateral bars
only distally (trellis imperfect) or throughout (trellis perfect). Calyptra mitrate, long-conic, or
long-cucullate, smooth, naked, sometimes clasping base of seta when young. Spores small or large, smooth to
lightly papillose. Genera
3, species 19 (3 genera and 15 species in the flora): North America, South
America, Eurasia, The
family Fontinalaceae is distinguished by aquatic or semi-aquatic habitat, long
axillary hairs, absence of stem central strand, lack
of stomata, odd exostome morphology, and the unusual trellised endostome. The
exostome teeth in the Fontinalaceae are linear with the outer and inner
plates equally high. The Fontinalaceae endostome is firm rather than
membranous because of relatively heavy secondary thickening. Secondary
deposition is extremely localized and occurs primarily on the outer surface at
and very near the wall pair sutures of the cells in the primary peristomial
layer (PPL). Deposition on the inner surface occurs only where the cell walls
of the inner peristomial layer (IPL) cells are contiguous to the secondary thickenings
deposited by the PPL cells. The Fontinalaceae endostome also differs in its
homogeneous structure; there is no division of the endostome into a basal
membrane and segment/cilia complex. As a result, the Fontinalaceae endostome,
or trellis, resembles a net consisting of 16 vertical filaments (each
corresponding to the endostomial segment of a typical endostome) that are
connected by nearly equidistant horizontal filaments. When
all the wall pair sutures in the PPL of a Fontinalaceae endostome are
secondarily thickened, an unbroken net called a perfect trellis is formed. An
imperfect trellis occurs when some wall pair sutures lack secondary
thickening. Whether the trellis is perfect or imperfect, all trellises have
16 vertical filaments. Thus, imperfect trellises are formed when some (usually
basal) horizontal wall pair sutures are unthickened. Furthermore, the
vertical filaments of the Fontinalaceae endostome have median lines on both
the front and back surfaces. In contrast, most moss endostomial segments have
a median line only on the outside surface. A somewhat similar endostomial
segment is seen in Funaria, Encalypta, Splachnum, Orthotrichum,
Mittenia, and Timmia. The combination, however, of a firm
endostome having a homogenous structure consisting of vertical filaments
connected by equidistant horizontal filaments and vertical filaments with
median lines on both surfaces is unique to the Fontinalaceae. All
three genera and most species of the Fontinalaceae are found in selected references Cardot, J. 1892. Monographie des
Fontinalacées. Mém. Soc. Sci. Nat. Cherbourg 28: 1--152. Welch, W. H. 1960. A
monograph of the Fontinalaceae. M. Nijhoff. 1. Leaves
ecostate or nearly ecostate................................................... 3.
Fontinalis, page XX 1. Leaves with long, single costa. 2. Leaves lanceolate or
linear-lanceolate, slenderly acuminate, acute or obtuse; calyptra covering
the capsule; seta 3--15 mm; capsules immersed, emergent or exserted......... 2. Dichelyma, page XX 2. Leaves
oblong-lanceolate, gradually acute; calyptra covering only the operculum; seta
0.7--1.5 mm; capsules immersed................................................. 1.
Brachelyma, page XX 3. FONTINALIS Hedwig, Sp. Musc. Frond.,
298. 1801 [Latin, fons, fontis, spring or fountain, alluding to
aquatic habitat] Plants glossy, yellow-red, yellowish green,
green, dark green, or brown. Primary stems creeping, secondary stems
trailing, freely and irregularly branched, stem and branch tips elongate,
slenderly and tightly attenuate or short and loose. Stems flaccid or rigid, in transverse section with epidermal
cells somewhat enlarged, subepidermal cells small, thick-walled; central
strand absent. Rhizoids on primary
stems and at base of secondary stems, from clusters of initials abaxially to
the leaf insertions, smooth, not or sparsely and irregularly branched. Leaves closely or distantly spaced,
imbricate to erect-appressed when dry, erect to erect-spreading when wet,
flaccid or firm, keeled and conduplicate, concave, or plane, ovate,
oblong-ovate, oblong-lanceolate, ovate-lanceolate, lanceolate, narrowly
lanceolate or linear, decurrent; apices acute, broadly acute or obtuse;
margins at times broadly reflexed proximally, plane, erect, or involute at
apex, entire proximally, entire to serrulate at apex; costa absent or nearly
absent; median cells linear-fusiform to long-rhomboidal, firm-walled, smooth;
alar cells enlarged, lax, bulging, subrectangular to oblong. Sexual condition dioicous; perigonia
and perichaetia lateral on short branches. Seta short, smooth, cells quadrate to subrectangular. Capsule immersed to slightly emergent,
ovoid to subcylindric; stomata absent; operculum conic to obtuse-conic;
annuli rudimentary, consisting of several rows of small, thick-walled,
horizontally elongated cells; peristome exostome teeth 16, linear, sometimes
united in pairs at the tips, outer surface smooth and lightly thickened,
inner surfaces heavily thickened, granulose, at times with obscure vertical
striae, inner trabeculae heavily thickened, endostome trellis perfect or
imperfect. Spores small or large,
smooth to lightly papillose, sometimes of two sizes due to developmental
abortions. Calyptra mitrate or
long-conic, smooth, naked. Species
13 (10 in the flora): North America, South America, Eurasia, Fontinalis
is characteristically aquatic
or semi-aquatic. Many species can be found in seasonally dry habitats, but all
are submerged in streams, lakes, ponds, pools, springs or seeps for a least
part of the year. The genus exhibits remarkable gametophytic variability,
perhaps due to the effects of seasonal wet/dry cycles as well as fluctuating
parameters associated with an aquatic habitat. Morphological variability is
high not only between populations but also within single collections of Fontinalis. The greatest amount of
morphological variability is in
species that normally are submerged only part of the year. Plants
of Fontinalis have creeping primary stems or stolons adnate to the
substrate and indeterminate in growth. Secondary stems are often sparsely,
irregularly branched and also indeterminate in growth; plants growing in
shallow, permanently flowing streams can be over three feet long. Fontinalis
is considered to have ecostate
leaves, but the leaves have a
multistratose basal region that has been interpreted as a rudimentary costa. Median
leaf cells are long and narrow; the alar cells are often enlarged and
inflated. Fontinalis has remarkably long axillary hairs; they can be over
900 \um. There are sporophytic character-complexes within Fontinalis that show meaningful
taxonomic patterns at the species level, but, since the genus is dioicous,
sporophytes are rarely encountered. Aquatic adaptations of sporophyte-associated
features in Fontinalis include perigonia with few antheridia (1--2 or
4--6); enlarged, well developed vaginula; immersed or emergent capsules; very
thick-walled capsules; lack of stomata; peristomes with endostomial trellises. The
taxonomy of Fontinalis is
complicated by the small number of useful gametophytic character states found
in the genus. In addition, because some important features of the genus are
macroscopic, proper identification requires having large, healthy collections.
Regrettably, success in keying out Fontinalis collections often
depends on prior experience with the genus. selected references Allen, B. 1983. On the costa of Fontinalis
(Musci). Lindbergia 9: 37--40. Allen, B. 1988. Studies on the genus Fontinalis
(Musci: Fontinalaceae). Brittonia 40: 180--187. Allen, B. 1991. A review of
the Fontinalis sullivantii complex (Musci: Fontinalaceae). Bryologist
94: 430--434. Allen, B. 1986--2008. Fontinalaceae Exsiccatae. Fasc. 1--6 (Nos.
1--174). 1. All or some
stem leaves keeled and conduplicate. 2.
Plants slender; leaves 2--4 mm. 3.
Stems flaccid; branch and stem tips flaccid, short-attenuate; median leaf
cells 4--8 \um wide; perichaetial leaves obtuse (gracilis-expression) 1. Fontinalis
antipyretica, in part 3.
Stems firm; branch and stem tips stiffly triangular, long-attenuate; median
leaf cells 10--12 \um wide; perichaetial leaves apiculate to cuspidate 5. Fontinalis neomexicana 2.
Plants medium-sized to robust; leaves 4--8 mm. 4.
All leaves keeled; keels more or less curved the entire leaf length 1. Fontinalis antipyretica, in part 4.
Stem leaves keeled or mostly keeled, branch leaves variously concave; keels
more or less straight distal to the basal curve. 5.
Branches with both keeled and concave leaves; branch and stem tips
triangular, swollen, and flaccid (oreganensis-expression)
1. Fontinalis antipyretica, in part 5.
Branches with only concave or tubular concave leaves; branch and stem tips
concave, firm but loosely foliate 3. Fontinalis
howellii 1. All leaves
concave or flat. 6.
All leaf apices with plane margins when wet. 7.
Stem and branch tips tightly long-attenuate. 8.
Plants with dimorphic leaves; stem leaves narrowly lanceolate to linear 9. Fontinalis sullivantii, in part 8.
Plants with stem and branch leaves intergrading in
size; stem leaves ovate-lanceolate, oblong-lanceolate, lanceolate, or
linear-lanceolate. 9.
Plants medium or large-sized, leaves flaccid, weakly concave or plane when
wet (vernalis-expression) 8.
Fontinalis sphagnifolia, in
part 9. Plants slender, leaves firm,
strongly concave when wet. 10.
Leaves linear-lanceolate, proximal margins not reflexed when dry (aestivalis-expression) 8. Fontinalis sphagnifolia, in part 10.
Leaves ovate-lanceolate to lanceolate, proximal margins usually reflexed when
dry 2. Fontinalis dalecarlica, in
part 7.
Stem and branch tips short-attenuate or loosely foliate. 11.
Leaves monomorphic; stem leaves ovate, ovate-lanceolate, oblong-lanceolate,
or narrowly lanceolate 4. Fontinalis
hypnoides 11.
Leaves strongly dimorphic; stem leaves lanceolate to linear. 12.
Stems flaccid; stem leaves 2--6(--9) mm, apices acute 9. Fontinalis sullivantii, in part 12.
Stems stiff; stem leaves 6--9 mm, apices obtuse 7. Fontinalis redfearnii 6. Leaf apices with erect to incurved
margins or a mixture of erect and plane margins when wet. 13.
Plants medium to large-sized, leaves 0.7--2 mm wide,
flaccid or firm, concave or plane when wet. 14.
Stem and branch tips tightly long-attenuate, leaves usually flaccid, weakly
concave with some plane leaves often present, leaf apices with margins when
wet involute or plane; median leaf cells long-rhomboidal, 60--120 \um (vernalis-expression) 8. Fontinalis
sphagnifolia, in part 14. Stem and branch tips usually loose
and turgid, at times tightly short-attenuate; leaves firm, concave
throughout, leaf apices with margins when wet consistently erect or involute;
median leaf cells linear-fusiform, 80--180 \um 6. Fontinalis novae‑angliae 13.
Plants slender, leaves 0.5--1.2 mm wide, firm, strongly concave when wet. 15.
Branch leaves linear-lanceolate (aestivalis-expression)
8. Fontinalis sphagnifolia, in part 15.
Branch leaves ovate-lanceolate to lanceolate. 16.
Leaves not dimorphic; leaf apices plane or occasionally erect when wet; dry
leaves usually flattened with narrowly reflexed margins; capsules immersed,
ellipsoid to oval; perichaetial leaves acute or apiculate 2. Fontinalis
dalecarlica, in part 16.
Leaves weakly dimorphic; leaf apices consistently erect to incurved when wet;
dry leaves without reflexed margins; capsules emergent, cylindric to
subcylindric; perichaetial leaves broadly rounded 10. Fontinalis welchiana 1.
Fontinalis antipyretica Hedwig, Sp. Musc. Frond.,
298. 1801 Hypnum antipyretica Necker; Pilotrichum antipyretica (Hedw.) Müller
Hal.; Fontinalis gigantea Sullivant; F. californica Sullivant; F.
antipyretica var. robusta Cardot;
F. antypyretica var. oreganensis Renauld & Cardot; F. mollis Müller Hal.; F. patula Cardot; F. allenii Cardot; F.
utahensis Cardot & Thériot in Thériot Plants medium to robust, to 40 cm, yellow-red,
yellowish green, green, dark green or brown. Axillary hairs to 820 \um, of 6--10 cells, basal cell red,
quadrate, distal cells hyaline, long-cylindric. Stems firm; stem and branch tips swollen, sharply angled. Leaves monomorphic, ovate,
oblong-ovate, ovate-lanceolate, or suborbicular, 2--8 mm, erect to
erect-spreading in 3 ranks when wet, imbricate to erect-appressed when dry, keeled
and folded lengthwise, variously curved along the keel from the insertion to
the apex; apices acute, broadly acute to rounded obtuse; margins occasionally
broadly reflexed proximally on one side, entire proximally ,
entire to serrulate at apices; median cells 100--150 x 12--18 \um. Perigonial leaves 1.2--1.4 mm. Perichaetial leaves 2--3 mm, oval to suborbicular, obtuse,
usually lacerated across the apex. Seta
0.2--0.3 mm. Capsule immersed to
slightly emergent, ovoid to subcylindric, 2--2.7 mm; operculum obtuse-conic,
0.7--1.5 mm; endostome trellis perfect. Calyptra
mitrate, 1.3--1.6 mm. Spores
14--20 \um. Rocks,
sticks, logs, roots in slow- or fast-moving streams, ponds, pools, ditches, swamps,
and floodplains, seasonally dry; low to high elevations (0--3300 m); Greenland;
St.Pierre and Miquelon; Alta., B.C., N. B., Nfld. and Labr., N.S., Ont.,
P.E.I., Que., Yukon; Alaska, Ariz, Calif., Colo., Conn., Del., Idaho, Ill., Maine,
Mass., Mich., Minn., Mont., Nev., N.H., N.J., N.Mex., N.Y., Ohio, Oreg., Pa.,
R.I., S.Dak., Utah, Vt., Wash., Wis., Wyo.; Europe; n and s Africa; Asia;
Atlantic Islands (Iceland). Fontinalis
antipyretica occurs in every conceivable habitat available to Fontinalis.
In its typical form the plants are medium-sized, flaccid when dry, have three-ranked, conduplicate-keeled leaves curved along the keel line, and swollen
stem and branch apices flaccid when dry. It is stenotypic in eastern Most
atypical western plants are variations on the oreganensis-expression: medium-sized plants; weakly keeled stem leaves
that are straight to slightly bend along the keel line; apices frequently
concave or plane and subobtuse or obtuse; branch leaves occasionally concave
or plane. The oreganensis-expression
is most common along the west coast, but also occurs sporadically throughout
the range of the species. Fontinalis
howellii and the oreganensis-expression
of F. antipyretica at times
intergrade; both have stem leaves with keels that are typically straight
above the basal curve and concave branch leaves. Fontinalis howellii differs from the oreganensis-form in having larger plants; an abrupt transition
from stem to branch leaf form; very long, slender branch leaves; and loosely
foliate, terete stem or branch tips that when dry lack the swollen, triangular,
appearance of the stem or branch tips in F. antipyretica. The
gracilis-expression is slender
(leaves 2--4 mm), loosely foliate, with lax to
flaccid stems and weakly keeled (often concave) leaves. It is similar to F. neomexicana in size and closely
approaches the loosely foliate expression of that species. Fontinalis neomexicana differs in
having firmer stems; more distinctly keeled leaves; narrower leaf cells (4--8
versus 10--12 \um wide); short-attenuate branch and stem tips; and apiculate
to cuspidate perichaetial leaves. Plants
of F. antipyretica found in swiftly
moving streams often have leaves completely split along the keel line, and
can be mistaken for F. hypnoides (see e.g., Crum & Anderson, Mosses
of North America 747). 2.
Fontinalis dalecarlica Bruch & Schimper in Bruch, Schimper & Gümbel,
Bryol. Eur. 5: 7. 1846 (fasc. 31) Fontinalis squamosa Hedwig var. dalecarlica (Bruch & Schimper)
Hartman Plants slender, to 100 cm, dark-green to
yellowish green, at times reddish. Axillary
hairs 250--300 \um, 5--8 cells, basal cell red, quadrate, distal cells
hyaline or reddish, long-cylindric. Stems
rigid; stem and branch tips long, slender and tightly attenuate; rhizoids
at base only. Leaves monomorphic,
ovate-lanceolate to lanceolate, 2--4 mm, imbricate to subimbricate wet, flat
with loosely reflexed margins dry, concave; apices acute or narrowly obtuse;
margins erect to incurved at base, plane or erect to incurved at apex,
serrulate or entire at apices; median cells 60--140 x 4--8 \um. Perigonial leaves 1--1.2
mm. Perichaetial leaves 2.8--3.2
mm, ovate to suboval, acute or apiculate, often lacerated across the apex. Seta 0.1--0.2 mm. Capsule immersed,
ellipsoid to oval, 1.5--2.5 mm; operculum obtuse-conic, 1 mm; endostome
trellis imperfect. Calyptra mitrate,
1.5--1.8 mm. Spores 20--50 \um. Rocks
and tree roots along or submerged in usually fast-moving streams,
occasionally in pools, and lakes; low to high elevations (0--1524 m); Greenland;
St.Pierre and Miquelon; N.B., Nfld and Labr., N.S., Ont., Que.; Ala., Conn.,
Ga., Ky., Md., Mass., N.H., N.J., N.Y., N.C., Pa., R.I., S.C., Tenn., Vt.,
Va., W.Va., Wis.; w Europe. Fontinalis
dalecarlica is a
slender species nearly always found in shallow, fast-moving streams. Wet
plants typically are smoothly imbricate with long attenuate branch and stem
tips; dry plants often have
flattened leaves with strongly reflexed margins. These features are characteristic,
but they appear to be accentuated by
and correlated with strong stream currents. When found in fast-moving streams,
the same features occur in F. neomexicana, F. novae-angliae, F. sphagnifolia, and F. hypnoides. Fontinalis dalecarlica has
monomorphic leaves that vary from strongly concave throughout to concave proximally
and plane distally. Apical leaf margins are usually plane,
but on strongly concave leaves they can be erect. The slender size and concave leaves of F.
dalecarlica distinguish it from all other members of the genus except F.
welchiana, F. sphagnifolia (aestivalis-expression), and rigid
forms of F. sullivantii. In
size, aspect, and branch leaf form F. welchiana and F. dalecarlica are
identical. Fontinalis welchiana,
however, typically occurs in seasonally dry streams, has weakly
dimorphic leaves (large subflaccid stem and small, rigid, strongly concave
branch leaves), and leaves with consistently erect to incurved apical leaf
margins. Fontinalis welchiana also differ from F. dalecarlica in
emergent, cylindric to subcylindric capsules and broadly rounded perichaetial
leaves. The aestivalis expression
of F. sphagnifolia is similar to F. dalecarlica in
having long-attenuate branch and stem tips as well as branch leaves that are
strongly concave when wet. It differs from F. dalecarlica in having long, linear-lanceolate leaves. Fontinalis sullivantii differs from F. dalecarlica in having strongly
dimorphic, linear-lanceolate stem and branch leaves with consistently plane
apical leaf margins. When dry the leaves in F. neomexicana collections from fast-moving streams often
resemble those of F. dalecarlica
because they are plane rather than keeled,
but when wet the leaves are strongly keeled. Large
plants of Fontinalis dalecarlica sometimes
approach those of F. novae-angliae in
size, and the two species are often confused. In addition to its larger size
(stem leaf width: 0.7--2 versus.0.5--1.2 mm), F. novae-angliae lacks
long-attenuate stem and branch tips, and its leaves have consistently erect
to incurved apical margins. It further
differs from F. dalecarlica in
having oval to suborbicular, obtuse perichaetial leaves; rhizoids at multiple
insertions along the stem; subcylindric to cylindric capsules; and smaller
spores (12--18 \um). All reports of F.
dalecarlica from 3. Fontinalis howellii Renauld & Cardot, Bot. Gaz.
(Crawfordsville) 13: 200. 1888 Fontinalis kindbergii Renauld & Cardot; F.
chrysophylla Cardot Plants medium to robust (rarely slender), to 35
cm, reddish, yellow-red, golden brown, to pale-green. Axillary hairs 600--750 \um, 6--10 cells, basal cell red,
quadrate to short-rectangular, distal cells hyaline, long-cylindric. Stems firm; stem and branch tips firm
and loosely foliate, terete, not angled. Leaves
strongly dimorphic. Stem leaves
ovate-lanceolate to lanceolate, 4--7 mm, erect to erect-spreading, loosely
ranked wet or dry, sharply or weakly keeled and folded lengthwise, keels
straight above the basal curve; apices acute to narrowly obtuse; margins plane,
at times erect to somewhat involute at apices, entire proximally, entire to
serrulate at apices; median cells 110--180 x 12--16 \um. Branch leaves lanceolate to linear-lanceolate, 3--6 mm,
erect-spreading when dry, spreading and sharply 3-ranked when wet, concave to
tubular-concave, not keeled; apices long-acuminate, acute to somewhat
rounded; margins involute. Perigonial
leaves 0.7 mm. Perichaetial leaves
2--3 mm, oval to suborbicular,
obtuse, usually lacerated across the apex. Seta 0.2--0.3 mm. Capsule
immersed to slightly emergent, oblong to subcylindric, 2.0--2.5 mm; operculum
conic, 1 mm; endostome trellis perfect. Calyptra
mitrate, 2 mm. Spores 12--16 \um. Logs,
sticks and shrubs in swamps, pools, and ponds, in streams, streamlets, or
springs, wet rocks in seeps, often seasonally dry; low to high elevations (3--1628
m); B.C.; Calif., Idaho, Mont., Oreg., Wash. Fontinalis howellii is typically a medium to robust
species found mostly in pools, ponds, or slow-moving water, often in
seasonally dry habitats, at times in association with Dichelyma. Wet
plants characteristically have spreading, sharply 3-ranked branch leaves;
when dry the branch and stem tips are firm, loosely foliate, and terete. The
typical form of F. howellii differs
from all other keeled species of Fontinalis
in having strongly heteromorphic leaves: stem leaves keeled-conduplicate;
branch leaves concave to tubular-concave. The change from stem to branch leaf
form is remarkably abrupt; transitional leaf forms are almost always absent. Also
distinctive are its stem leaves with straight keels after the basal curve,
and linear-lanceolate to lanceolate branch leaves. Plants that grow
permanently submerged (chrysophylla-expression)
differ from the typical form in their smaller size and in having mostly concave
stem leaves. The
oreganensis-expression of Fontinalis antipyretica and F. howellii at times intergrade; both
have stem leaves with keels that are typically straight above the basal curve
and concave branch leaves. The oreganensis-expression
differs from F. howellii in often
having smaller plants; some transitional leaves between the typical stem and
branch leaf forms; shorter, broader branch leaves; and triangular branch and
stem tips that when dry have a swollen aspect very different from the firm, loosely
foliate, branch and stem tips in F. howellii. All collections referred to F. howellii from eastern North
America and recent reports from 4. Fontinalis hypnoides
Hartman, Handb. Skand. Fl. (ed. 4) 434. 1843 Fontinalis duriaei Schimper; F. nitida Lindberg & Arnott; F.
tenella Cardot; F. dalecarlica var.
macounii; F. mac-millanii Cardot; F.
holzingeri Cardot in Holzinger;
F. obscura Cardot in Holzinger;
F. umbachii Cardot; F. subcarcinata Cardot; F. nitida var.
angustiretis Cardot ex W.
H. Welch. in Grout Plants slender to medium-sized, to 30 cm,
green, yellowish green, or pale green. Axillary
hairs to 800 \um, 6--10 cells, basal cell red, quadrate, distal cells
hyaline or reddish, long-cylindric. Stems
flaccid, stem and branch tips loose, occasionally tightly attenuate. Leaves typically widely spaced,
spreading, ovate, ovate-lanceolate, lanceolate, narrowly lanceolate to
linear-lanceolate, 3--7 mm, flaccid dry, plane or subconcave at base, apices
broadly acuminate, acute or subobtuse, occasionally twisted at the apices;
margins plane or erect proximally, plane at apices, entire to serrulate at
apex; median cells 80--150 x 8--15 \um. Perigonial
leaves 0.6--0.8 mm. Perichaetial
leaves 1.8--2 mm, oval to suborbicular, obtuse, usually lacerated across
the apex. Seta 0.25--0.3 mm. Capsule immersed to slightly
emergent, oval, suboval or oblong, 1.5--2.5 mm; operculum obtuse-conic, 0.8--1.2
mm; endostome trellis perfect. Calyptra
mitrate, 1.2--1.5 mm. Spores
13--20 \um. Rocks,
boulders, base of trees, roots, sticks in streams, swamps, lakes, or ponds; low
to high elevations (6--2800 m); Alta., B.C., Man., N.B., N.W.T., N.S., Ont.,
Que., Sask., Yukon; Ala., Alaska, Ariz., Ark., Calif., Colo., Conn., Idaho, Ill.,
Ind., Iowa, Kans., Ky., Maine, Md., Mass., Mich., Minn., Mo., Mont., Nev.,
N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Oreg., Pa., S.Dak., Tex., Utah,
Vt., W.Va., Wis., Wyo; Europe; n Asia. Fontinalis
hypnoides is separated
from all other Fontinalis species
by medium size and monomorphic leaves consistently plane at the apices. It is
typically found in slow-moving to sluggish streams and ponds. Critical
features of the species in this habitat include flaccid stems with
well-spaced leaves that are plane or subconcave only at the very base and have
enlarged, distinctly differentiated alar cells. Plants from fast-moving
streams differ from the typical form in having stiffer stems with closely
spaced leaves that are subconcave to concave in the lower half and sometimes
have reflexed margins when dry. Leaf shape in F. hypnoides is extremely variable: ovate to ovate-lanceolate (duriaei-expression); lanceolate to
narrowly lanceolate (hypnoides-expression);
linear-lanceolate (tenella-expression).
Extreme expressions are sometimes recognized at the species level; but so
many collections have leaves with intermediate forms that the group is best
treated as an intergrading series of expressions. A particularly distinctive
expression often found in the northern Flaccid forms of F. hypnoides can be
difficult to separate from F. sphagnifolia (vernalis-expression)
which is similar in size, has monomorphic leaves, and at times flaccid stems with mostly plane leaves.
Plants of F. sphagnifolia differ from those of F. hypnoides in
having tight, long-attenuate branch and stem tips,
and a mixture of leaf apices with plane or erect to incurved margins. Stiff
forms of F. hypnoides can resemble F. novae-angliae, but that species has
leaf apices with erect to incurved margins. Both stiff and flaccid forms of F. hypnoides are sometimes confused
with F. sullivantii which also has
stiff and flaccid forms, well-spaced, plane to subconcave leaves, and
consistently plane leaf apices. Fontinalis
sullivantii always has linear-lanceolate stem leaves and most of its
forms differ from F. hypnoides in
their considerably smaller size. All collections of F. sullivantii differ from F.
hypnoides in having strongly dimorphic stem and branch leaves. Fontinalis redfearnii and F. hypnoides (tenella-expression) are similar in aspect and have linear-lanceolate
leaves with plane apical margins. Fontinalis
redfearnii, however, has strongly dimorphic leaves and longer stem (6--9 mm)
leaves. 5.
Fontinalis neomexicana Sullivant
& Lesquereux, Musci Bor.-Amer. 47 [no. 224b]. 1856 [1857] Fontinalis mercediana Lesquereux; F. maritima Müller Hal.; F.
columbica Cardot Plants slender to medium, to 50 cm,
pale-green, green, yellowish green, or golden brown.
Axillary hairs 300--450 \um, 5--7
cells, basal cell red, quadrate, distal cells hyaline, long-cylindric. Stems firm; branch and stem tips
imbricate, long-attenuate, sharply 3-angled. Leaves monomorphic, oblong-lanceolate to ovate-lanceolate, 2--4(--5)
mm, erect, erect-spreading in 3 ranks when wet, erect-spreading to spreading when
dry, keeled and folded lengthwise, keel straight or moderately curved above
basal curve, occasionally strongly curved; apices acute, occasionally
subobtuse; margins at times loosely reflexed when dry, plane when wet, entire
throughout, occasionally serrulate at apices; median cells 120--180 x 4--8 \um.
Perigonial leaves 1.3--1.5 mm. Perichaetial leaves 3--3.5 mm, oval to suborbicular, abruptly
apiculate to cuspidate, at times lacerated across apex. Seta 0.2--0.3 mm. Capsule
immersed to slightly emergent, ovoid to oblong, 2--2.5 mm; operculum
obtuse-conic, 0.6--0.9 mm; endostome trellis perfect. Calyptra mitrate, 2 mm. Spores
15--25 \um. Rocks
and boulders, rarely sticks, swift or occasionally slow streams; low to high
elevations (0--3353 m); Alta, B.C.; Fontinalis
neomexicana is a
slender species with monomorphic, keeled leaves, very narrow leaf cells, and
mostly entire leaf margins. Leaf keels above the basal curve vary from nearly
straight to moderately curved. Apiculate to
cuspidate perichaetial leaves are a diagnostic feature of the species. Fontinalis
neomexicana is often
found in shallow, fast-moving streams. In this habitat wet plants are
typically triangularly imbricate with firm, long attenuate, sharply 3-angled
branch and stem tips; dry plants often
have erect-spreading to spreading, distinctly 3-ranked leaves with
loosely reflexed margins. These features appear to be accentuated by and are correlated
with strong stream currents. Dry plants with loosely reflexed leaf margins are
also associated with fast-moving stream habitats in F. dalecarlica, F. novae-angliae, F. sphagnifolia, and F. hypnoides. In slow-moving
streams, F. neomexicana has loosely
foliate stems and somewhat lax, shortly attenuate branch and stem tips. The
imbricate foliate form of F.
neomexicana is similar in size
and aspect to F. dalecarlica;
both species have attenuate branch and stem tips. They are sometimes confused
because the dry leaves of F.
neomexicana can appear to be concave rather than keeled. Wet plants of F. neomexicana differ from F. dalecarlica in having keeled leaves
and angled rather than terete branch and stem tips. Slender forms of F. antipyretica (gracilis-expression) closely approach the loosely foliate form of
F. neomexicana; the slender forms
of F. antipyretica, however, have
lax to flaccid stems; weakly keeled (often concave) leaves; broader leaf
cells (10--12 vs. 4--8 \um wide), short, swollen branch and stem tips; and
obtuse perichaetial leaves. 6.
Fontinalis novae-angliae Sullivant, Musci Hepat. U. S. (repr.)
p. 104. 1856 Fontinalis lescurii Sullivant var. cymbifolia Plants medium-sized to robust, green,
yellowish green, reddish green to brownish. Axillary hairs 500--650 \um, 5--8 cells, basal cell red,
quadrate, distal cells hyaline or reddish, long-cylindric. Stems stiff to firm; stem and branch
tips swollen, not or slightly attenuate. Leaves
monomorphic, ovate, ovate-lanceolate, or oblong-lanceolate, 2.5--5.5 mm, erect
to imbricate wet, concave to tubular-concave; apices acute to subtruncate;
margins erect or plane at base, erect to incurved at apex, serrulate or
occasionally entire at apices; median cells 80--180 x 8--15 \um. Perigonial leaves 1--1.2 mm. Perichaetial leaves 2.4--2.8 mm, oval
to suborbicular, obtuse, usually lacerated across apex. Seta 0.1--0.3 mm. Capsule
immersed to slightly emergent, subcylindric to cylindric, 1.5--2.5 mm;
operculum obtuse-conic, 1 mm; endostome trellis imperfect to subperfect. Calyptra mitrate, 1.2--1.5 mm. Spores 12--18 \um. Rocks,
boulders, and roots in ponds, lakes, streams and rivers, at times in summer
dry streams; low to medium elevations (3--600 m); N.B., Nfld. and Labr.
(Nfld), N.S., Ont., Que.; Ala., Ark., Conn., Del., D.C., Fla., Ga., Ky., La.,
Maine, Md., Mass., Miss., N.H., N.J., N.Y., N.C., Okla., Pa., S.C., Tenn.,
Tex., Vt., Va., W.Va. Fontinalis
novae-angliae is a
medium-sized to robust species typically found in slow-moving streams, lakes
or ponds. It has swollen to slightly attenuate branch and stem tips, and monomorphic,
ovate-lanceolate to oblong-lanceolate leaves with consistently erect to
involute apical margins. Dry plants typically have concave leaves, but plants from fast-moving streams sometimes
have flat leaves with narrowly reflexed margins when dry. There are two
distinct expressions of F. novae-angliae. The type-expression has somewhat lax plants with well-spaced, broad,
moderately concave leaves. In the cymbifolium-expression the plants
are stiff with closely spaced, narrow, tubular-concave leaves. Fontinalis howelli is the only other species in the genus
with tubular-concave leaves. In that species, however, the branch leaves are
tubular-concave but the stem leaves are keeled-conduplicate. Fontinalis novae-angliae
has been chronically confused with F. sphagnifolia. The two species
are often similar in size and leaf shape, but F. sphagnifolia is a morphologically
complex species with two remarkably distinct forms: the vernalis-expression and the aestivalis-expression.
Both forms of F. sphagnifolia differ from F. novae-angliae in
having long, tightly attenuate stem and branch tips. The vernalis-expression of F. sphagnifolia further differs in
flaccid leaves with apical margins that are sometimes erect to incurved and
other times plane. The aestivalis-expression
has firm, strongly concave leaves similar in form to those of the cymbifolium-expression of F. novae-angliae, but differs in
having distinctly smaller, slender plants with linear-lanceolate leaves. Fontinalis
dalecarlica and F.
novae-angliae have often been confused because they both have firm,
concave leaves. But F. dalecarlica is
a slender species with narrow leaves (stem
leaves 0.5--1.2 versus 0.7--2 mm wide) that usually have plane apical leaf
margins when wet and reflexed proximal leaf margins when dry. In addition the
stem and branch tips of F. dalecarlica are typically long-attenuate;
the perichaetial leaves are acute or apiculate; the capsules are immersed,
ellipsoid to oval; and the spores are larger (20--50 versus 12--15 \um). 7.
Fontinalis redfearnii B.
H. Allen, Bryologist 94: 201. 1991 Plants robust, to 15 cm, dark-green to
yellowish green. Axillary hairs 740--918 \um, 8--12 cells, basal 1--2 cells reddish brown, short-rectangular,
distal cells hyaline, becoming red with age, long-cylindric. Stems
rigid; stem tips loosely foliate. Stem
and branch leaves dimorphic, differing greatly in size. Stem leaves long, narrowly lanceolate
to linear, 6--9 mm, distantly spaced, erect-patent dry, stiffly patent to
spreading wet, firm, concave at base, plane; apices broadly obtuse; margins
plane at apex, entire proximally, entire to serrulate at apex; median cells
80--160 x 8--12 \um. Branch leaves
3--4 mm, concave, apices acute to obtuse. Sporophytes unknown. Base
of Taxodium, rocks; medium
elevation (243 m); Fontinalis
redfearnii is separated from other members of Fontinalis by its robust size; distantly spaced leaves; loosely
foliate stem tips; strongly dimorphic branch and stem leaves; and narrowly
lanceolate to linear-lanceolate stem leaves that have broadly obtuse,
consistently plane apices. Three other Fontinalis
species with wide-spaced, narrowly lanceolate leaves with plane apices
can be confused with F. redfearnii:
F. hypnoides (tenella-expression), F.
sphagnifolia (vernalis-expression), and F. sullivantii. Fontinalis
hypnoides (tenella-expression)
differs from F. redfearnii in
having flaccid to somewhat stiff stems and smaller (5--7 mm), monomorphic
leaves. F. sphagnifolia (vernalis-expression) differs most clearly from F. redfearnii in having tight,
long-attenuate branch and stem tips. In addition, it has flaccid stems,
weakly dimorphic leaves, and a mixture of leaf apices with plane or erect to
incurved margins. Fontinalis
sullivantii is the only other Fontinalis
species with strongly dimorphic leaves. Although F. sullivantii typically differs from F. redfearnii in having much smaller plants and flaccid stems, there
are robust, stiff forms of the species that closely approach F. redfearnii. All forms of F. sullivantii, however, differ from F. redfearnii in narrower leaves with
acute apices. 8.
Fontinalis sphagnifolia (Müller Hal.) Wijk & Margadant,
Taxon 11: 221. 1962 Pilotrichum sphagnifolium Müller Hal., Syn. Musc. Frond. 2: 150.
1850; Fontinalis biformis Sullivant in A. Gray; F. lescurii Sullivant in A. Gray; F.
eatoni Sullivant ex Sullivant
& Lesquereux; F. lescurii var. ramosior Sullivant; F.
frostii Sullivant in Cardot;
F. novae-angliae var. cardotii Renauld in Cardot; F. cardotii (Renauld
in Cardot) Cardot; F. novae-angliae var. heterophylla Cardot in Nichols; F. novae-angliae var. latifolia
Cardot in Nichols; F. novae-angliae var. groutii W. H. Welch in Grout Plants slender, medium-sized, or robust, to
60 cm, green, yellowish green, reddish green or brownish. Axillary hairs 500--650 \um, 5--8
cells, basal cell red, quadrate, distal cells hyaline or reddish, long-cylindric.
Stems flaccid or rigid, stem and
branch tips long, tightly attenuate. Stem
leaves extremely variable in form, closely or distantly spaced, erect,
erect-spreading, or spreading, ovate, ovate-lanceolate, oblong-lanceolate, or
lanceolate, 3--6 mm, firm or lax, concave throughout to flaccid, weakly
concave or plane, acute to obtuse; margins erect or somewhat plane at base,
erect-incurved or plane distally, erect-incurved or plane, serrulate or
entire at apex; median cells 60--120 x 6--15 \um. Branch leaves similar to stem leaves except smaller, or closely
spaced, erect to erect-spreading, narrowly lanceolate or linear-lanceolate, 2--3.5
mm, firm, concave to subtubular, long-acuminate; margins involute throughout,
erect to involute, serrulate at apex. Perigonial
leaves 0.8--1.1 mm. Perichaetial
leaves 2.4--3.4 mm, oval to suborbicular, obtuse, usually lacerated
across apex. Seta 0.1--0.2 mm. Capsule immersed to slightly
emergent, subcylindric to cylindric, 2--2.5 mm; operculum conic, 1 mm;
endostome trellis imperfect, cilia connected only in distal half in a
lattice. Calyptra mitrate, 1.3--1.5
mm. Spores 16--18 \um, lightly
papillose. Rocks,
roots, stumps, logs, in streams that are often seasonally dry; low to high
elevations (2--1005 m); Greenland; Alta., Man., N.B., Nfld. and Labr.
(Nfld.), N.S., Ont., Que.; Ala., Ark., Conn., Del., D.C., Fla., Ga., Ill., Ind.,
Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., Mo., N.H., N.J.,
N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., Tenn., Vt., Va., W.Va., Wis. Fontinalis
sphagnifolia has long,
tightly attenuate branch and stem tips and non-keeled leaves with often relatively
short, long-rhomboidal median leaf cells. In all other features, however, it
is remarkably variable. Typically the species is found in shallow streams and
pools that are seasonally dry. The regular seasonal alternation between
aquatic and terrestrial environments produces plants with two distinctly
different leaf forms. Leaves formed after a prolonged dry period (vernalis-expression) or in sluggish
water are medium-sized, loosely flaccid, ovate-lanceolate, plane to weakly
concave, and have plane or erect to incurved apical margins. Leaves formed
late in the aquatic period (aestivalis-expression)
or in faster-moving streams are smaller in size, stiff, linear-lanceolate,
strongly concave and have consistently erect to incurved apical margins. There
are many collections that have a combination of vernalis-expression and aestivalis-expression
leaves; whether or not this combination is the result of leaves formed during
the transition from aquatic to terrestrial environments is unknown. Most of
these intermediate collections fall within the lescurii-expression of F.
sphagnifolia. The leaves in this expression are small and
linear-lanceolate, but flaccid and have a mixture of plane and erect apical
margins. When well developed the vernalis-expression
of F. sphagnifolia has stem leaf apices with quadrate to
short-rhomboidal leaf cells, and this has been erroneously considered an
important feature of the species. Apical leaf cell shape within Fontinalis
appears to be positively associated with leaf apex shape: the broader the
leaf apex the shorter and broader the apical leaf cells. Fontinalis
novae-angliae,
the duriaei-expression of F. hypnoides, and F. redfearnii are similar to the vernalis-expression of F.
sphagnifolia in size and aspect.
Fontinalis novae-angliae
differs from the vernalis-expression
of F. sphagnifolia in having short,
often tumid branch and stem apices
and firm leaves that are consistently concave with incurved to involute
apical margins. The duriaei-expression of F. hypnoides is
similar to the vernalis-expression
of F. sphagnifolia in having weakly concave to plane leaves; it
differs in having leaves with consistently plane apical leaf margins and
longer median leaf cells. Fontinalis
redfearnii differs from F.
sphagnifolia (vernalis-expression)
in having loosely foliate stem tips. Fontinalis dalecarlica has been confused with the aestivalis-expression of F.
sphagnifolia because of similar
size, long, slender, tightly attenuate branch and stem tips, and strongly
concave leaves. Fontinalis dalecarlica
differs in ovate-lanceolate to lanceolate rather than narrowly lanceolate or
linear-lanceolate leaves that are flat with loosely reflexed margins when
dry. In addition, the apical leaf margins in F. dalecarlica vary from plane to erect. Large expressions of F. sullivantii can be very similar in
size and aspect to the lescurii-expression
of F. sphagnifolia, but that
species has consistently plane apical leaf margins. Although F.
sphagnifolia has generally been unrecognized in eastern 9. Fontinalis sullivantii Lindberg,
Oefvers. Förh Finska Vetensk.-Soc.. 12: 77. 1869
[1870] Fontinalis disticha Hooker & Wilson in Drummond; F. disticha Hooker & Wilson
var. tenuior Sullivant; F. lescurii Sullivant var. gracilescens
Sullivant; F. dichelymoides Lindberg;
F. filiformis Sullivant & Lesquereux in Austin; F. flaccida Renauld
& Cardot; F. renauldii Cardot; F. langloisii Cardot; F.
filiformis var. tenuifolia Cardot; F. missourica Cardot; F. microdonta Renauld ex Cardot Plants slender to medium-sized, to 25 cm,
dark green, green, yellowish green, or pale green. Axillary hairs 480--600 \um, 6--10 cells,
basal cell red, quadrate, distal cells hyaline or reddish, long-cylindric. Stems rigid or flaccid; stem and
branch tips short or long attenuate. Stem
and branch leaves dimorphic, differing greatly in size. Stem leaves narrowly lanceolate to
linear, 2--6(--9) mm, widely spaced, erect-spreading to spreading, flaccid or
firm, subconcave at base, plane, concave, or tubular above; apices acuminate,
acute to subobtuse; margins plane at apex, entire proximally, entire to
serrulate at apex; median cells 80--150 x 4--8 \um. Branch leaves 2--4 mm. Perigonial
leaves 1.0--1.2 mm. Perichaetial
leaves 2--3 mm, oval to suborbicular, obtuse, usually lacerated across
apex. Seta 0.25--0.30 mm. Capsule immersed to slightly emergent
narrowly cylindric, 1.7--2.5 mm; operculum conic, 0.8--1 mm; endostome
trellis imperfect. 12--20 \um. Calyptra
mitrate, 1.0--1.3 mm. Spores
12--20 \um, lightly papillose. Rocks,
roots, tree bases, logs, sticks, and twigs, in streams, sloughs, swamps, or
pools; low to medium elevations (0--580 m.); N.S.; Ala., Ark., Conn., Del.,
Fla, Ga., Ill., Ky., La., Maine, Md., Mass., Miss., Mo., N.J., N.H., N.Y.,
N.C., Ohio, Okla., Pa., R.I., S.C., Tenn., Tex., Va.; n Europe. Fontinalis
sullivantii is separated from other members of Fontinalis by its typically slender size and narrowly
lanceolate to linear-lanceolate, strongly dimorphic branch and stem leaves
that have consistently plane apical margins. Plants growing in pools or slow-moving
streams have flaccid stems and limp, plane leaves. Those found in fast-moving
streams have rigid stems and stiff, subtubular branch leaves. In addition
plants of F. sullivantii can vary
considerably in branching pattern (regular or sparse) and stem leaf length (4
or more versus 3 or less mm). Fontinalis sullivantii has four major expressions (see Allen
1991) that have somewhat discrete ranges. Recognizing them at the species
level is unwarranted because of the many intermediate morphology collections
that link the expressions together. Three other slender species of Fontinalis with
variously plane or concave leaves can be confused with F. sullivantii:
F. dalecarlica, F. welchiana, and F. sphagnifolia (aestivalis-expression).
Fontinalis dalecarlica resembles rigid forms of F. sullivantii, but
it has monomorphic leaves that are strongly concave when wet, and flattened when dry with narrowly reflexed margins. Fontinalis welchiana and the aestivalis-expression
of F. sphagnifolia often have
weakly dimorphic leaves, and they can have the same flaccid aspect as F. sullivantii. They differ from F. sullivantii in having strongly
concave branch leaves with erect to incurved apical margins. Dichelyma
capillaceum is superficially similar to F. sullivantii in its
slender aspect, widely spaced branches, and long, narrow leaves. It differs from
F. sullivantii in having keeled
leaves with long, single costae. Fontinalis redfearnii is the only other Fontinalis species with strongly dimorphic leaves. It differs
from most collections of F. sullivantii
in its much larger size and stiffer stems, but there are robust, stiff forms
of F. sullivantii that closely
approach F. redfearnii in size. Fontinalis redfearnii, however, differ
from all forms of F. sullivantii in
having broadly obtuse stem leaf apices. 10.
Fontinalis welchiana B. H. Allen, Brittonia 40: 181. 1988 Plants slender, to 20 cm, green, yellow-green,
or reddish yellow. Axillary hairs 450--530
\um, 5--10 cells, basal cells red, quadrate, distal cells hyaline or reddish,
long-cylindric. Stems stiff; stem
and branch tips shortly attenuate. Leaves
weakly dimorphic or monomorphic. Stem
leaves oblong-lanceolate, 2.5--4 mm, erect wet, erect-spreading dry, moderately
concave; apices acute to obtuse, serrulate or entire; margins erect to
incurved at apex; median cells 50--80 x 5--8 \um. Branch leaves ovate-lanceolate to lanceolate, 1.5--2 mm, strongly
concave; apices acute, serrulate; margins incurved at apex; median cells 40--65
x 7 \um. Perigonial leaves 0.7--0.8
mm. Perichaetial leaves 2.6--3 mm,
ovate to suboval, broadly rounded, often lacerated across apex. Seta 0.2 mm. Capsule emergent, cylindric to subcylindric, 2--3 mm; operculum
conic, 0.5 mm; endostome trellis nearly perfect, latticed ½--3/4 their length.
Calyptra mitrate, 1.5 mm. Spores 20--40 \um. Rocks,
ledges, and roots, in streams or seasonally dry pools, also constantly
submerged in rivers and spring runs; low to medium elevations (152--580 m);
Ark., Ill., Mo., Okla. Fontinalis
welchiana is a slender
species of shallow streams that are seasonally dry; but also permanently
submerged in rivers or deep spring runs. Plants from the two habitats have
distinctly different aspects. Those exposed to seasonal drying have a soft
aspect with weakly dimorphic leaves that are often plane to subconcave below
when dry. Plants that are permanently submerged are stiff with monomorphic, long-lanceolate
leaves that are strongly concave wet or dry. Both forms have smoothly imbricate to attenuate branch and stem tips, and leaves
with erect to incurved apical margins. Fontinalis
sullivantii, F. sphagnifolia (aestivalis-expression), and
F. dalecarlica are slender species with variably plane or concave leaves that
can be confused with F. welchiana. In aspect, stiff forms of F.
sullivantii are especially difficult to distinguish from the permanently
submerged expression of F. welchiana. But F. sullivantii lacks attenuate branch and stem tips, and has
strongly dimorphic leaves with plane apical margins. Both forms of F.
welchiana can be confused with F.
sphagnifolia (aestivalis-expression) and F. dalecarlica because they all have attenuate
branch and stem tips and concave leaves that when dry can have erect to
incurved apical leaf margins. Fontinalis
sphagnifolia (aestivalis expression)
differs in having very long-attenuate branch and stem tips, and lanceolate to
linear-lanceolate branch leaves. Fontinalis
dalecarlica and F. welchiana are
remarkably alike in aspect and branch leaf form; F. dalecarlica differs in having monomorphic leaves that when wet have plane or erect to
incurved apical margins, and when dry are usually flattened with narrowly reflexed
margins. Fontinalis dalecarlica also differ from F. welchiana in
having immersed, ellipsoid to oval capsules, and acute or apiculate perichaetial
leaves. |
