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BFNA Title: Philonotis |
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Philonotis -
Bartramiaceae V. PHILONOTIS
Bridel, Bryol. Univ. 2:15. 1827 * [Greek philos,
lover; notis, moisture] Dana Griffin, III Plants small to robust, hygrophilic, glaucous
to whitish green or yellowish green, in dense to lax tufts. Stems
0.5--10(--16) cm, erect, simple, forked or with a subfloral whorl of
branches, more or less tomentose proximally; epidermis of large, thin walled,
hyaline cells, often with exterior wall collapsed. Leaves in many
rows, rarely in 5 distinct rows, 1- stratose, erect-spreading or somewhat
secund when dry, erect-spreading to spreading when moist, broadly to narrowly
lanceolate, acute to acuminate, rarely obtuse; margins plane or revolute,
serrulate usually throughout, teeth single or paired; costa subpercurrent to
excurrent, smooth or rough abaxially; distal laminal cells subquadrate to
oblong or linear, prorulose at distal or proximal ends on both surfaces,
rarely smooth or with a centric papilla; basal cells usually more lax than
distal cells; alar cells not or slightly differentiated. Specialized
asexual reproduction lacking or small deciduous brood branches in axils
of distal leaves. Sexual condition dioicous, less frequently autoicous
or rarely synoicous; perigonia gemmiform or discoid; perigonial leaves
scarcely or clearly distinct from stem leaves; perichaetia terminal or
lateral by innovations; perichaetial leaves scarcely distinct from stem leaves.
Seta solitary, usually elongate, straight or flexuose, rarely curved. Capsule
globose to ovoid, erect to horizontal to inclined, furrowed or rarely
irregularly wrinkled, mouth oblique; annulus none; operculum conic convex,
blunt to mammillate or bluntly apiculate; peristome double or rarely lacking;
teeth lanceolate, dark red to reddish brown, densely and finely papillose,
trabeculate, typically with rounded to ovoid thickenings on interior surface;
endostome yellowish to pale brown; basal membrane well developed; segments
keeled, vertically striate papillose; cilia 1--3, well developed or
rudimentary. Spores spherical to reniform, densely and usually
coarsely papillose. Species 178
(11 in the flora): worldwide except Antarctica.. SELECTED REFERENCES Crum,
H.A. and L.E. Anderson. 1981. Mosses of Eastern North America, vol. 1. New
York. Florschűtz, P. A. 1964. The Mosses of Suriname, part 1. Leiden.
Flowers, S. 1935. Philonotis. In:
A. J. Grout, ed., Moss Flora of North America. Newfane, Vermont. Ireland, R.
R. 1982. Moss Flora of the Maritime Provinces, Publications in Botany 13,
National Museums of Canada. Ottawa. Nyholm, E. 1954. Illustrated Moss Flora
of Fennoscandia II. Musci. Gleerup. Lund. Zales, W. M. 1973. A Taxonomic
Revision of the Genus Philonotis
for North America north of Mexico, Ph.D. dissertation (unpubl.). University
of British Columbia. Vancouver. Philonotis will be recognized by the 1-stratose
leaves and the subfloral whorl of branches on fertile plants. While leaf
cells are typically prorulose, sterile stems, especially when subject to
periodic inundation, may bear leaves with smooth cells. 1. Synoicous;
seta curved; mature capsules wrinkled to more or less smooth; plants small.
. . 1. Philonotis cernua 1. Autoicous
or dioicous; seta straight or flexuose; mature capsules furrowed;
plants small to robust. 2.
Leaf cells with a centric papilla . . 2. Philonotis
yezoana 2.
Leaf cells prorulose from one or both ends or, occasionally, prorulae obscure
or absent. 3.
Costa subpercurrent; some or most leaves broadly acute or rounded-obtuse;
laminal cells smooth or prorulose at distal end; plants small and delicate .
. . 3. Philonotis gracillima 3.
Costa percurrent to long-excurrent; leaves acute to acuminate (rarely with
some leaves obtuse and costa subpercurrent, in which case laminal cells
prorulose at proximal end); plants small to robust. 4.
Leaf cells prorulose at proximal end or, occasionally, at either end on
adaxial side. 5.
Leaves spirally imbricate, seriate; costa coarsely prorulose abaxially from
tip to base; Greenland . . . . . .. 9. Philonotis
seriata 5.
Leaves not or rarely spirally imbricate, not seriate; costa smooth or only
weakly prorulose abaxially, distally. 6.
Plants robust; costa 300--600 µm wide at base; cells near costa at widest
part of leaf 48--100 µm; perigonial leaves acute; Greenland . . . . . . . 10.
Philonotis calcarea 6.
Plants small to robust; costa to 320 µm wide at base; cells near costa at
widest part of leaf 24-40 µm; perigonial leaves obtuse or, if acute, then
stem leaves with long-excurrent costa
. . . . . .. . . . . . . 11. Philonotis
fontana 4.
Leaf cells prorulose at distal end throughout or, occasionally, at both ends
on abaxial side. 7.
Leaf cells elongate (5--20:1), in longitudinal rows. 8.
Leaves longly triangular-lanceolate, straight or falcate, flat; cells long
and narrow (9--20:1); prorulae pointed, projecting at extreme distal end of
cell; autoicous . . . . 4. Philonotis
longiseta 8.
Leaves triangular to slightly ovate-lanceolate, keeled; cells rectangular
(less than 9:1); prorulae rounded, near distal end of cells; dioicous . . . .
. . . 5. Philonotis marchica 7.
Leaf cells quadrate (1:1) to rectangular (1--4:1), in less obvious
longitudinal rows. 9.
Prorulae conspicuous, numerous; leaf margins strongly revolute, scabrous,
marginal teeth sharp; costa long-excurrent; Alabama and Florida . . . . .6. Philonotis
sphaerocarpa 9.
Prorulae neither conspicuous nor numerous, sometimes obscure and few; margins
revolute, not scabrous, marginal teeth more or less blunt; costa variable. 10.
Leaf margins singly serrulate; costa long-excurrent; Pacific coast, northern
Rocky Mountains . . . . 7. Philonotis capillaris 10.
Leaf margins doubly serrulate; costa percurrent to excurrent; southeastern
United States . . . . .8. Philonotis uncinata 1. Philonotis cernua (Wilson) D. G.
Griffin & W. R. Buck, Bryologist 92: 376. 1989 Glyphocarpa cernua
Wilson, Hook. J. Bot. 3: 383. 1841; Bartramidula
carolinae Flowers Plants glaucous or yellowish, delicate, in
loose to dense tufts. Stems erect, 0.5--(-2) cm. Leaves laxly
erect-appressed, occasionally somewhat secund when dry, erect-spreading when
moist, lanceolate, acute to acuminate, 0.5--1 mm; margins plane to weakly
revolute, serrulate distally; costa percurrent, rough abaxially; distal
laminal cells rectangular, 20--30 × 4--6
µm, prorulose at one or both ends, firm-walled; basal cells similar but wider
(6--8 µm wide). Sexual condition synoicous; perichaetial leaves
scarcely differentiated. Seta curved to flexuose, 0.5--1.5 mm, smooth.
Capsule erect to pendulous, globose with short tapering neck, 0.8--1.5
mm, leptodermous, smooth to irregularly wrinkled when dry; operculum low
convex, occasionally with a low, blunt beak; peristome lacking but with a low
membrane inserted proximally the mouth. Spores reniform, 36--44 µm. Capsules
mature Mar.--Oct. Wet acidic rocks in mountains of southeastern United
States; 1200--1500 m; NC, Tenn.; Mexico; Central America; South America;
Europe (U.K.); possibly Africa. This is a
diminutive species that can be identified by the synoicous sexual state, the
globose, smooth to irregularly wrinkled capsules and the lack of a peristome. 2. Philonotis yezoana Bescherelle &
Cardot in J. Cardot, Bull. Soc. Bot. Geneve, ser. 2, 1: 123. 1909 Didymodon mollis
Schimper, Syn. Eur., ed. 2, p. 167. 1876 not Philonotis mollis (Dozy
& Molkenboer) Mitten 1859 nor Venturi 1882 Plants dark green, rufous-tomentose
proximally, in dense tufts or intermixed with other bryophytes. Stems
1--3 cm, erect, simple or sparingly branched. Leaves laxly
erect-appressed when dry, often somewhat incurved, erect-spreading when
moist, ovate-lanceolate, sometimes acuminate, 0.8--1.2 mm; margins narrowly
revolute near base, plane distally, serrulate throughout; costa subpercurrent
to excurrent, smooth abaxially; laminal cells firm-walled, with a single
centric papilla on both sides (papillae more prominent on adaxial side);
distal cells rectangular, 15--26 × 6--8
µm; median cells short-rectangular, 15--20 × 8--10
µm; basal cells similar to median cells but subquadrate to quadrate. Specialized
asexual reproduction by slender brood branches in leaf axils. Sexual
condition dioicous. [Perigonia discoid. Seta 2.5--4 mm. Capsule
spherical, 2--3 mm. Spores 19-24 µm]. Rocky cliffs
or steep slopes, wet or dry sites; 0--2700 m; B.C., Nfld., Ont.; se Alaska,
Calif., Mont., Wash., Vt.; Asia (Japan; Korea). The dark green
color and centrally papillose leaf cells make Philonotis yezoana one
of the easiest to recognize of the flora species. Characters of
sporophyte-bearing plants (Japan) are from W.M. Zales (1973). 3. Philonotis gracillima Ångstrom, Oefv. K. Sv. Vet. Ak. Foerh. 33(4):
17. 1876 Plants small, delicate, bright green, in
dense to loose tufts. Stems 0.5--1 cm; erect, simple to sparingly
branched. Leaves distant to laxly imbricate, erect to incurved when
dry, erect-spreading to spreading when moist, 0.3--1 mm, ovate oblong to
ligulate or ovate-lanceolate to ovate, apex broadly acute to rounded-obtuse;
margins revolute throughout or plane distally and revolute proximally,
bluntly serrulate by paired teeth; costa extending 7/8 of lamina or
subpercurrent, rough abaxially distally; laminal cells lax, pellucid, thin
walled, distal cells subquadrate to oblong rhomboidal, 20--30 × 10--12 µm, obscurely to clearly prorulose at distal ends,
basal cells rectangular, 25--35 × 10--15
µm. Specialized asexual reproduction by occasional short brood
branches in axils of distal leaves. Sexual condition dioicous. [Perigonia gemmiform, subtended by
1--5 branches, often appearing lateral. Seta ca. 20 mm. Capsule
1--1.5 mm, furrowed when dry; peristome as for genus. Spores reddish
brown, 20--26 µm, papillose]. Moist, rocky,
usually limey soil or in rock crevices; 0--1500 m; Ala., Fla., La., Miss.,
N.C., Kans., Okla., Tex; Mexico; Central America; West Indies; South America. Sporophyte traits
are as given by P.A. Florschütz (1964), who treated this species as a variety
of Philonotis uncinata. Philonotis gracillima is distinguished in the floral range by the
combination of a broad leaf apex (both broadly acute and rounded-obtuse
apices may occur on the same plant), relatively short costa and the lax,
pellucid leaf cells. Specimens of similar morphology have been collected from
many parts of the Old World tropics under an older name, P. hastata (Duby) Wijk
& Margadant. Further study is required to resolve this possible synonymy. 4. Philonotis longiseta (Michaux) E.
Britton, Bryol. 14: 44. 1911 Bartramia longiseta
Michaux., Fl. Bor.-Amer. 2: 301. 1803 Plants bright green in loose to dense tufts,
rufous-tomentose proximally. Stems 1--3 cm, simple or forked. Leaves
crowded, laxly erect to erect, sometimes secund when dry, erect-spreading
when moist, 1--2.5 mm, narrowly triangular-lanceolate, acuminate; margins
narrowly revolute, serrulate nearly throughout; costa excurrent; laminal
cells prorulose throughout, prorulae projecting forward over distal ends of
cells; distal cells linear, 30--65 × 5--7
µm, basal cells shorter and broader. Sexual condition autoicous
; perigonia gemmiform, lateral and just proximally perichaetia. Seta
1.5--3.5 cm. Capsule 1.8--2 mm. Spores reniform, 26--33 µm,
papillose. Capsules
mature Feb.--Dec. Soil or rock, often on wet embankments; Ala., Ark., Fla.,
Ga., Iowa, Kans., La., Miss., Mo., N.Y., Ohio, Okla., Penn., S.C., Tex., Va.;
Mexico; West Indies (Puerto Rico); Central America (Guatemala, Costa Rica):
South America (Venezuela). Philonotis longiseta
can be recognized by the flat, narrowly triangular leaves with forward
projecting prorulae at the distal ends of the laminal cells throughout the
leaf. 5. Philonotis marchica (Hedwig) Bridel,
Bryol. Univ. 2:23. 1827 Mnium marchicum
Hedw., Sp. Musc. 196. 1801; Philonotis
muehlenbergii (Schwaegrichen)
Bridel, Bryol. Univ. 2: 22. 6. 1827 Plants slender, in yellowish or bright green
tufts, brown-tomentose proximally. Stems erect, simple or forked, 1--6
cm. Leaves 1--2.3 mm, erect-spreading when dry, somewhat divergent
when moist, occasionally spiraled, triangular-lanceolate, acuminate, keeled;
margins plane or sometimes narrowly revolute, serrulate nearly to base, teeth
single; costa percurrent to long-excurrent; distal laminal cells linear
oblong, 20--30 × 4--8 µm, prorulose , prorulae rounded,
near the distal ends of cells or occasionally at both ends, basal cells
oblong, 18-45 × 6--15 µm. Specialized asexual
reproduction by propagulae occasionally borne in leaf axils. Sexual
condition dioicous; perigonia discoid. Seta 1.5--4 cm. Capsule
1--2.5 mm. Spores 20--30 µm, papillose. Capsules
mature Mar.--Sept. Rocks and soil in wet places, roadsides, springs; 35--3000
m; Alta, B.C., Man., N.S., Que.; Ala., Ariz., Ark., Colo., Fla., Ga., Idaho,
Ill., Ind., Iowa, Kans., Mich., Minn., Miss., Mo., Nebr., N.C., N.H., N.
Mex., N.Y., Ohio, Okla., Oreg., Pa., Tenn., Tex., Utah, Va., Vt., W. Va.,
Wis., Wyo.; Mexico; Central America; South America (Colombia); Europe; Asia;
n Africa; Atlantic Islands (Macaronesia). Philonotis marchica
is recognized by the usually plane, singly serrulate leaf margins and the
laminal cells bearing rounded prorulae near the distal ends. Philonotis capillaris, a species mainly of Pacific coastal habitats, shares
with P. marchica the singly serrulated leaf margin; however, P. capillaris,
in contrast to P. marchica, has a more or less
decumbent, flaccid habit, more distant, widespread leaves, shorter and
broader distal laminal cells and obscure prorulae in the basal half of the
leaf. 6. Philonotis sphaerocarpa (Hedwig)
Bridel, Bryol. Univ. 2: 25. 1827 [as “sphaericarpa”] Mnium sphaerocarpon
Hedwig, Sp. Musc. 197. 1801
Plants relatively robust, yellowish green, in
dense tufts, rufous-tomentose proximally. Stems 1--3 (--6) cm, simple
or forked. Leaves erect, straight or somewhat homomallous when dry,
erect-spreading when moist, 1.2--2 mm, lanceolate acuminate; margins
sharply serrulate, teeth paired but
often appearing in groups of 3 or 4 because of strongly revolute margins;
costa long-excurrent, scabrous abaxially distally; laminal cells strongly
prorulose at distal ends, distal and median cells narrowly oblong, 18--35 × 3--5 µm, basal cells rectangular, 12--25 × 6--8 µm. Sexual condition dioicous. [Seta
1.5--2.5 cm. Capsule 1.5--2 mm. Spores subspherical to
reniform, 23--26 µm, papillose.] Open soils,
road banks; 10--100 m; Ala., Fla. La.; Mexico; West Indies; Central America;
n South America. Philonotis sphaerocarpa
is identified by the long-excurrent, scabrous costa, strongly revolute leaf
margin, and leaf cells bearing prominent prorulae especially on adaxial side.
While not uncommon in the tropical portion of its range, the species is
relatively rare in the flora area. 7. Philonotis capillaris C. Hartman, Skand. Fl., ed. 2. 10:
40. 1871 Plants delicate, scattered or in loose,
yellowish to light green tufts, tomentose proximally. Stems usually
simple, weakly erect to procumbent, 1--3 cm. Leaves erect when dry,
somewhat distant, erect-spreading when moist, lanceolate from an ovate base,
acuminate, 0.5--1.5 mm; margins plane to narrowly revolute, serrulate nearly
to base, teeth single, projecting from distal ends of marginal cells; costa
excurrent, rough abaxially distally; distal and median laminal cells oblong,
10--30 × 5--7 µm, prorulose at distal ends,
basal cells shorter and broader, short-rectangular to quadrate, prorulose at
distal or sometimes at proximal ends. Sexual condition dioicous;
perigonia discoid. Seta 2--3 cm. Capsule 1.3--2 mm. Spores
subreniform, 20--26 µm, papillose. Capsules
mature Apr.-- Aug. Moist sandy soil or humus, sometimes on rock ledges, often
in shady habitats; 50--2300 m; B.C.; Alaska, Calif., Idaho, Oreg., Wash.;
Europe; sw Asia. Philonotis capillaris
is distinguished by the unpaired teeth of the leaf margin that project from
the distal ends of the cells and by the relatively short laminal cells with
prorulae at the distal ends. In the flora area, the species is of restricted
distribution, basically limited to Pacific coastal habitats from California
to Alaska with incursions eastward to the western slopes of the Idaho
Rockies. It is associated with an oceanic climate. 8. Philonotis uncinata (Schwaegrichen)
Bridel, Bryol. Univ. 2: 22. 1827 Bartramia uncinata
Schwaegrichen, Suppl. Sp. Musc. 1(2): 60. 1816; Philonotis glaucescens
(Hornschuch) Brotherus, Bih. K. Svensk. Vet. Ak. Handl. 21 Afd. 3(3): 27.
1895 Plants small, in dense to lax, yellowish to
yellowish green tufts, tomentose proximally. Stems simple, erect to
inclined, straight or curved, 0.5--1.5(--3) cm. Leaves
triangular-lanceolate, 0.7--1.3 mm, erect and straight or curved and
homomallous when dry, erect-spreading when moist; margins plane proximally,
narrowly revolute distally, serrulate nearly to base, teeth paired distally,
usually single proximally; costa percurrent to short-excurrent, rough
abaxially distally; laminal cells prorulose at distal ends, distal cells
narrowly oblong, 20--50 × 5--8 µm, basal cells wider, quadrate
to rectangular, 10--65 × 8--12 µm. Specialized asexual
reproduction by brood branches in the axils of distal leaves. Sexual
condition dioicous; perigonia gemmiform. Seta erect, 1.3--3 cm. Capsule
ovoid, 1.5--2 mm. Spores subspherical to reniform, 23--26 µm,
papillose. Capsules
mature Feb.--Mar. Rocks and soil in open habitats; 0--30 m; Ala., Fla., Ga.,
Ky., La., Miss., S.C., Tex.; Mexico; West Indies; Central America; n and c
South America; Pacific Islands. This species,
restricted in the flora area largely to states bordering the Gulf of Mexico,
is recognized by the diminutive habit, the percurrent to short-excurrent
costa, doubly toothed leaf margin and laminal cells prorulose at distal ends.
Philonotis glaucescens, treated as a synonym here, has been recognized
elsewhere as P. uncinata var. glaucescens (Hornschuch) Florschütz (P. A. Florschütz 1964) or as
P. glaucescens (Hornschuch) Brotherus (Crum and Anderson 1981). The
features on which "P. glaucescens"
is recognized (whether as a variety or as a species) - straight leaves and a
percurrent costa - would seem to fall well within the universe of variation
displayed by P. uncinata, the oldest name available
for this group of related forms. 9. Philonotis seriata Mitten, J. Linn.
Soc. Bot. Suppl. 1:63. 1859 Philonotis fontana
var. seriata (Mitten) Kindberg,
Bih. K. Svensk. Vet. Ak. Handl. 7(9): 255. 1883 Plants robust, in greenish to yellowish
tufts. Stems erect, simple, 3--12 cm. Leaves erect to
erect-spreading and imbricate when dry, erect-spreading when moist, plicate,
straight or falcate, seriate in 5 spiraled rows, ovate-lanceolate, 1.5--2 mm;
margins plane or narrowly revolute, serrulate to denticulate nearly to base,
teeth paired or simple; costa robust, percurrent to excurrent, rough abaxially
throughout; laminal cells prorulose at proximal ends, prorulae mostly on
adaxial side, distal cells narrowly oblong to linear, 25--45 × 4--6 µm, basal cells rectangular to elongate-hexagonal,
15--30 × 7--10 µm. Sexual condition
dioicous; perigonia discoid. Seta erect, 3--5 (--7) cm. Capsule
horizontal, furrowed when dry, 2--3 mm. Spores subreniform, 20--22 µm,
papillose. Capsules
mature June--Sept. Acid seeps and springs, high mountains or at high northern
latitudes; 10--3000 m; s Greenland; Europe; Asia (India; Iran, e, n and w
Russia; n Africa). Philonotis seriata
is an Old World species that enters the flora area only in southern
Greenland. It is allied with P. fontana, including having obtuse inner
perigonial leaves, but differs in the seriate stem leaves and the costa that
is scabrous on the back throughout. According to S. Flowers (1935), authentic
P. seriata does not occur on the North American continent. 10. Philonotis calcarea (Bruch, Schimper & W. Gümbel) Schimper, Coroll. 86. 1856 Bartramia calcarea
Bruch, Schimper & W. Gümbel, Bryol. Eur. 4: 49. 325. 1842 Plants robust, in bright green to yellowish
green tufts, brownish tomentose proximally. Stems erect, simple, 4--8
cm. Leaves on sterile and female stems appressed, usually secund when
dry, erect-spreading to falcate spreading when moist, 1.3--3 mm, plicate,
ovate-lanceolate to lanceolate; margins revolute with paired teeth on mature
leaves; costa very stout (to 600 µm wide at base), subpercurrent to
short-excurrent, somewhat scabrous abaxially distally; laminal cells
prorulose at proximal ends; distal laminal cells linear, 20--40 × 3--5 µm, basal cells shorter and broader, juxtacostal
cells at widest part of leaf lax, pellucid and quite large (48--100 × 10--30 µm). Sexual condition dioicous; perigonia
discoid. Seta to 4 cm. Capsule 2--3.5 mm. Spores
subreniform, 22--25 µm, papillose. A member of
the Philonotis fontana complex, this essentially Old World species enters the
flora area only in southern Greenland. Its distinguishing features include
the falcate secund leaves, very stout costa, and lax, pellucid and quite
large juxtacostal cells. 11. Philonotis fontana (Hedwig)
Bridel, Bryol. Univ. 2:18. 1827 Mnium fontanum Hedwig, Sp. Musc. 105.
1801 Plants small to robust, in light to dark
green or yellowish green tufts, mats or sods, occasionally reddish or
glaucous, typically reddish brown tomentose proximally. Stems 1--16
cm, erect, simple, irregularly branched or with a subfloral whorl of
innovations. Leaves stiffly erect to erect or erect-spreading, less
commonly catenulate, sometimes falcate or falcate secund, rarely distalmost
leaves spiraled around stem, lanceolate to broadly ovate-lanceolate or
ovate-subulate, plane, bi- or pluriplicate, 0.6--3 mm, gradually to abruptly
narrowed to acumen, apex acute to acuminate, occasionally obtuse; margins
revolute, serrulate throughout, teeth paired and appearing 2-fid due to their
apposing position from contiguous cells, occasionally with margins plane and
teeth unpaired; costa short- to long-excurrent (often subpercurrent in obtuse
leaves), to 320 µm wide at base; laminal cells prorulose at proximal ends on
abaxial side and at proximal and distal ends on adaxial side, distal cells
linear to oblong-linear, 15--40 × 3--5
µm, basal cells more lax, rectangular to oblong-hexagonal, up to 24--40 × 7--10 µm near costa. Sexual condition dioicous;
perigonia discoid. Seta straight, 2--5(--7) cm. Capsule globose
to ovoid, 1--3.5 mm. Spores ovoid to reniform, 18--30 µm, papillose. Capsules
mature throughout the year, with the season of maturation influenced by,
among other things, latitude and elevation. Rock or soil, often in seepy,
open habitats; 0--3500 m; Greenland; widespread in North America from Alaska
to Mexico; c and w Europe; Asia; c and n Africa. Philonotis
fontana has a
Holarctic distribution with limited penetration into the montane tropics of
both Eastern and Western Hemispheres. Even given its membership in a seepage
community, where morphological plasticity is not uncommon, the extent of variation
in this species is excessive. Many variants have been recognized but with
little firm evidence to support the majority of them. E. Nyholm (1954) was
convinced that only through a series of cultivation, cytological and genetic
studies could the immense variability within this polymorphic complex be
properly evaluated. W. M. Zales (1973) was able to show by a comparison of
cultured and field-derived plants which of the morphological characters were
relatively stable and which were subject to environmental influence
("ecophenic characters"). His treatment of this complex, with minor
deviation, is followed here. The core characters for the complex are laminal
cells prorulose at proximal ends on the abaxial side, juxtacostal cells near
the leaf base 24--40 µm, teeth of the leaf margin typically paired and
appearing 2-fid, and costa up to 320 µm wide at the leaf base. 1. Plants
small; leaves stiffly erect, not plicate; capsules 1--2 mm . . . . . Philonotis fontana var. pumila 1. Plants
small to robust; leaves erect to spreading or catenulate, occasionally
appressed, plane, bi- or pluriplicate; capsules 2--3.5 mm. 2.
Plants robust; stems to 16 cm; leaves distant, spreading, catenulate,
pluriplicate, distalmost leaves spiraled . . . .Philonotis fontana var. americana 2.
Plants small to robust; stems to 10 cm; leaves imbricate, erect to spreading,
not catenulate, sometimes appressed, plane or 2-plicate, distalmost leaves
not spiraled . Philonotis fontana
var. fontana 11a. Philonotis fontana var.
fontana Plants small to robust. Stems 2--20
cm. Leaves imbricate, erect to spreading, sometimes appressed, not
catenulate, plane or with a single pair of plicae near leaf base. Capsule
2--3 mm. Capsules
mature throughout the year. Rock or soil in seepy, typically open habitats;
0--3500 m; Greenland; Alta., B.C., Man., N.B., Nfld., N.S., N.W.T., Nunavut,
Ont., Que., Sask., Yukon; Ala, Alaska, Ariz., Calif., Colo., Fla., Ga.,
Idaho, Ill., Kans., Maine, Mich., Mo., Mont., Nebr., Nev., N.Mex., N.Y.,
N.C., Pa., S.C., S.Dak., Tenn., Tex., Utah, Vt., Va., Wash., Wyo.; Mexico; c
and w Europe; Asia; e and n Africa. While the
breadth of variation in the typical variety is immense, crucial characters
include the typically robust habit, the imbricate, laxly erect, straight,
falcate or secund leaves. Plants that are subject to prolonged periods of
submersion may develop bizarre morphology. 11b. Philonotis fontana var.
americana (Dismier) H. A.
Crum, Bryologist 72: 244. 1969 Philonotis
americana Dismier,
Bull. Soc. Bot. France 10. mem. 17: 35. 1910 Plants robust to quite robust. Stems
5--16 cm. Leaves distant, widespread, catenulate, sometimes falcate,
distalmost leaves spiraled around stem, broadly ovate lanceolate,
pluriplicate. Capsule 2.5--3.5 mm. Capsules
mature June--Sept. Seeps, exposed slopes or mountain meadows; 0--2940 m;
Alta, B.C., N.B., Ont., Que.; Alaska, Calif., Colo., Idaho, N.Y., Oreg.,
Utah, Wash. This variety,
which includes the largest plants in the genus in North America, can be
recognized on the basis of the robust habit, distant, catenulate leaves with
the distalmost leaves spiraled around the stem. llc. Philonotis fontana var.
pumila (Turner) Bridel,
Bryol. Univ. 2:20. 1827 Bartramia
fontana var. pumila Turner, Musc. Hib. 107.
1804 Plants small. Stems 1--6 cm. Leaves
stiffly erect, plane. Capsule 1--2 mm. Capsules
mature June --Aug. Seepage slopes and along creeks, often over clay or silt,
sometimes intermixed with other bryophytes; 0--3250 m.; Alta., B.C., Man.,
N.S., N.W.T., Nunavut, Ont., Que., Yukon; Alaska, Calif., Colo., Minn.,
Mont., Oreg., Tenn., Wash., Wyo.; c and n Europe; c and sw Asia; Atlantic
Islands (Iceland). This
diminutive variety typically grows in dense mats or sods, the stems tightly
interlaced with tomentum. The stiffly erect leaves that are neither
catenulate nor spiraled aid in its identification. The range is
Arctic-alpine. This variety is a characteristic member of the bog community
throughout the Arctic tundra and taiga. |