BFNA Title: Philonotis
Author: D. Griffin, III  
Date: November 26, 2003
Edit Level: R Brum+
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

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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.