PLANTS PLEUROCARPOUS

Plants pleurocarpous; stems creeping, spreading, forming mats, or pendent, frondose or dendroid, often forming tufts, from a creeping primary stem; sporophytes lateral on stems; peristome double or variously reduced, rarely absent.


Section 6. Plants pleurocarpous. Leaves 2-ranked, strongly folded throughout.


1. Leaves to 1.5 mm long; apex mostly mucronate and reflexed; laminal cells entire, not porose Catagoniaceae
1. Leaves 3 mm or more long; apices truncate or erect apiculate; laminal cells porose Phyllogoniaceae



Section 7. Plants pleurocarpous. Leaves ecostate.

1. Plants aquatic, attached to rocks in streams or epiphytic in inundated lowlands; capsules immersed Fontinalaceae
1. Plants terrestrial, on soil, rocks, logs or epiphytic, if aquatic then alar cells inflated; capsules mostly exserted 2
2. Leaves broadly ovate, acute to piliferous, narrowly bordered, border differentiated in colour, often dark red; laminal cells appearing finely pluripapillose, papillae minute and numerous over and between cells Rhacocarpaceae
2. Leaves variously shaped, acuminate, acute, or obtuse rounded; laminal cells smooth or if papillose, papillae single or few over cell lumina 3
3. Alar cells inflated, oval to oblong, mostly thick-walled, porose or not, often dark yellow or golden-red; laminal cells smooth or papillose, papillae over cell lumina Sematophyllaceae p. p.
3. Alar cells absent or if present, then not inflated or oval, mostly subquadrate or oblate and quadrate- to short oblong-rounded 4
4. Leaf apex narrowly long acuminate Leucomiaceae
4. Leaf apex broadly acute or if acuminate then laminal cells rhomboidal to fusiform-rounded and thick-walled 5
5. Leaves short ovate-lanceolate, to 0.7 mm long; laminal cells on back projecting at distal angles; alar cells equally distributed on either side of leaf base, mostly quadrate; plants forming mats Myriniaceae p.p.
5. Leaves variously shaped, ovate-lanceolate, ovate to oblong, mostly greater than 1 mm long; laminal cells smooth, or occasionally on back projecting at distal angles; alar cells lacking or equally or unequally distributed on either side of leaf base, quadrate to oblong; plants forming mats, dendroid or pendent 6
6. Stems complanate foliate, lateral leaves slightly asymmetric or not 7
6. Stems terete foliate 9
7. Leaf margins finely bifid toothed; gemmae clustered beneath stems or on specialised terminal branches; epiphytic or epiphyllous Pilotrichaceae p.p.
7. Leaf margins entire to subentire; gemmae absent or if present on leaf tips; epiphytic on base trunk of trees, logs or soil 8
8. Alar cells well differentiated, unequally distributed with cells on one side more numerous; upper laminal cells linear; gemmae absent; moist to wet lowlands Stereophyllaceae p.p.
8. Alar cells undifferentiated; upper laminal cells large, elongate-hexagonal; gemmae few on leaf tips and distal margins; montane forests Hookeriaceae
9. Leaf margins plane distally; laminal cells obliquely arranged above, transversely toward base or not; median and upper cells rhomboidal to fusiform-rounded, porose or not 10
9. Leaf margins strongly incurved distally; laminal cells longitudinally arranged, linear, mostly porose 11
10. Leaves broadly lanceolate and plicate, or ovate and smooth, short acuminate; capsules exserted Leucodontaceae
10. Leaves broadly ovate to suborbiculate, smooth, broadly acute, or if acuminate then tip hyaline; capsules immersed Erpodiaceae
11. Plants dendroid, or subpendent and rather stiff; gemmae commonly present in leaf axils Pteryobryaceae p.p.

11. Plants pendent or forming loose, spreading mats; gemmae absent

Meteoriaceae p.p.


Section 8. Plants pleurocarpous. Leaves costate; costae short and forked or double and elongate (often 1/2 or more than lamina length).


1. Leaves strongly cordate-auriculate; paraphyllia present, several branched Hylocomiaceae p.p.
1. Leaves not cordate-auriculate, if auriculate only weakly so; paraphyllia absent or if present then linear, unbranched 2.
2. Leaf costa double and elongate, often 1/2 or more the lamina length Pilotrichaceae p.p.
2. Leaf costa mostly short and forked, mostly less than 1/2 the lamina length 3.
3. Leaves complanate 4.
3. Leaves teretely foliate, falcate-secund or not 9.
4. Leaf margins bordered by few to several rows of narrow cells, inner cells large, rhomboidal to hexagonal; costa often indistinct or distally forked from a short single costa; seta papillose and ciliate distally Daltoniaceae p.p.
4. Leaf margins lacking a border, occasionally cells of lamina progressively narrowed toward margin 5
5. Leaves ovate-oblong to oblong-lingulate, often undulate; apices mostly broadly acute to obtuse; capsules immersed to shortly exserted Neckeraceae p.p.
5. Leaves mostly ovate to ovate-lanceolate, smooth; apices mostly acute to long acuminate; capsules exserted 6.
6. Alar cells numerous, extending along leaf margin 1/5-1/3 7.
6. Alar cells few to several, rarely extending more than a few cells along margin; capsules erect to horizontal 8.
7. Laminal cells smooth; capsules erect Entodontaceae p.p.
7. Laminal cells papillose at cell angles; capsules inclined Pterigynandraceae
8. Distal branches flagellate and readily deciduous; seta distally roughened to papillose; urn spinose Symphyodontaceae
8. Distal branches not flagellate, if so not readily deciduous; seta smooth; capsules lacking ornamentation 8
9. Leaves weakly serrulate to subentire distally, short to long decurrent; capsules suberect to horizontal, cylindrical Plagiotheciaceae
9. Leaves often serrulate to serrate throughout, not or weakly decurrent; capsules ovoid and horizontal to subpendent or cylindrical and erect Hypnaceae p.p.
10. Leaves falcate-secund or branches and stems distally curved or curled 11.
10. Leaves mostly erect to spreading, not falcate-secund, branches and stems not curved 12.
11. Alar cells quadrate to short rectangular, moderately thick-walled, concolourous or yellow to golden, or cells inflated and hyaline, thin-walled; exothecial cell walls of urn mostly evenly thickened; operculum conic-apiculate to -short rostrate Hypnaceae p.p.
11. Alar cells inflated, thick-walled, often golden-red, supra-alar cells often present, subquadrate; costa weakly short and forked or absent;exothecial cell walls often collenchymatous; operculum long rostrate Sematophyllaceae p.p.
12. Stems either dark red with leaves concave and ovate with alar cells subquadrate and thick-walled (often dark red), or stems wiry with leaves concave and ovate, weakly plicate, costa separated at base; alar cells enlarged and thin-walled Hylocomiaceae p.p.
12. Plants lacking the combination of characters given above Amblystegiaceae p.p.


Section 9. Plants pleurocarpous. Leaves costate; costa single. Leaves strongly dimorphic, upper or lower leaves on stem smaller and differing in shape from larger lateral leaves.


1. Leaves on upper side of stem smaller; leaf margins lacking a border; costa of lateral leaves stoutly short excurrent Racopilaceae
1. Leaves on lower side of stem smaller; leaf margins bordered or at least differentiated; costa of lateral leaves ending well below apex or weakly short excurrent Hypopterygiaceae


[Sections 10-11. Plants pleurocarpous. Leaves monomorphic, usually differing only in that branch leaves are somewhat smaller, differing in shape or not, or that lateral leaves asymmetric and median leaves symmetric, or primary and secondary stem leaves differentiated; costa single.]


 Section 10. Plants pleurocarpous. Leaves costate; costa single; laminal cells mammillose or papillose.


1. Laminal cells pluripapillose (cells with a single papillae mixed with some cells occasionally 2-3-papillose keyed out here also) 2
1. Laminal cells uniformly unipapillose or mammillose 6
2. Papillae rather numerous, over cell lumen and side walls Meteoriaceae p.p.
2. Papillae few, confined to cell lumen 3
3. Paraphyllia present on stems; leaves of stem and branch differentiated, the former ovate or triangular and acuminate, the latter ovate and acute or obtuse-rounded; stems regularly 1-3 pinnately branched; leaf apices ending in one or more papillae Thuidiaceae p.p.
3. Paraphyllia absent; leaves of secondary stems and branches differing mostly in size; stems irregularly branched; leaf apices not papillose 4
4. Upper laminal cells isodiametric, mostly obscurely hexagonal Anomodontaceae p.p.
4. Upper laminal cells elongate, linear to oblong-linear 5
5. Secondary stems rather rigidly erect, solitary or few branched; leaf margins strongly recurved Pterobryaceae p.p.
5. Secondary stem soft or rigid, not erect, several to numerously branched; leaf margins not recurved, or if so only weakly so at base Meteoriaceae p.p.
6. Stems differentiated between creeping primary stems and erect, subpendent or frondose / dendroid secondary stems 7
6. Stems not differentiated between primary and secondary stems, plants mostly forming mats or dense tufts 12
7. Creeping stems densely tomentose, leaves mostly obscurely hidden among tomentum; elongate branches (appearing as secondary stems) erect and often numerous; seta elongate, on terminal branches; calyptra mitrate-campanulate, often deeply lobed and plicate Orthotrichaceae p.p.
7. Creeping stems radiculose beneath, or naked; secondary stems erect to suberect, pendent or frondose to irregularly branched; seta lateral on stems 8
8. Upper laminal cells with papillae projecting at distal angles 9
8. Upper laminal cells with papillae over cell lumen 10
9. Plants not stipate, mostly irregularly branched; gemmae absent; seta very short, capsules immersed, sporophytes often several to rather numerous along one side of stems
Cryphaeaceae p.p.
9. Plants mostly stipate, often frondose; gemmae often present; seta elongate, capsules exserted Pteryobryaceae p.p.
10. Leaf margins forming a border of somewhat elongate, smooth cells Meteoriaceae p.p.
10. Leaf margins lacking a border, cells similar to intralaminal cells 11
11. Plants small, frondose; leaves ovate, smooth; upper laminal cells rhombic, walls entire Neckeraceae p.p.
11. Plants medium sized to large, irregularly branched to subdendroid; leaves ovate-lanceolate, plicate; upper laminal cells rhomboidal to linear-rhomboidal, stellate Prionodontaceae
12. Stem leaf margins strongly ciliate; laminal cell papillae strongly curved Anomodontaceae p.p.
12. Stem leaf margins serrulate to serrate, not ciliate; laminal cell papillae erect or projecting 13
13. Plants forming tufts; leaves falcate-secund; some cells with papillae strongly projecting at distal angles on back of leaf Hylocomiaceae p.p.
13. Plants forming mats; leaves not falcate-secund; papillae over cell lumen 14

14. Leaves loosely complanate, lateral leaves somewhat asymmetric, oblong acute-rounded; alar cells unequally distributed, more numerous on one side of costa

Stereophyllaceae p.p.

14. Leaves not complanate, ovate acuminate; alar cells equally distributed on either side of costa

 Leskeaceae p. p.



Section 11. Plants pleurocarpous. Leaves costate; costa single; laminal cells smooth.


1. Laminal cells elongate, mostly 8 or more times longer than wide 2
1. Laminal cells isodiametric to short oval or rhomboidal, mostly 5 or less times longer than wide 6
2. Plants pendent, dendroid or frondose, from a creeping primary stem (occasionally forming loose mats on banks or logs); stem and branch leaves often dimorphic or polymorphic in repeated series 3
2. Plants spreading or creeping, stems not differentiated between primary and secondary; stem and branch leaves differing only by degree of size and often by width 4
3. Plants mostly erect and frondose or dendroid, if pendent then leaves in 5-spiral rows; numerous filamentous pseudoparaphyllia present Pteryobryaceae p.p.
3. Plants pendent or occasionally forming loose spreading mats; leaves not arranged in 5-spiral rows; filamentous pseudoparaphyllia absent Meteoriaceae p.p.
4. Leaf alar cells asymmetrically distributed, more numerous on one side of costa Stereophyllaceae p.p.
4. Leaf alar cells, if present, equally distributed on either side of costa 5
5. Plants mostly of wet habitats; leaves smooth; alar cells usually present, either thin-walled, enlarged and rounded or if quadrate to short rectangular then cells small and few; capsules horizontal, strongly asymmetric and curved Amblystegiaceae p.p.
5. Plants of moist or semi-dry habitats, if aquatic then leaves broadly ovate, leaf margins serrulate throughout, upper laminal cells ca. 2:1; leaves plicate or smooth; alar cells quadrate to short rectangular, often rather numerous; capsules inclined to erect, weakly asymmetric Brachytheciaceae p.p.
6. Stems differentiated between creeping primary stems and erect to suberect secondary stems 7
6. Stems creeping or spreading, occasionally subascending, not differentiated between primary and secondary stems 14
7. Secondary stems distinctly stipate, subdendroid to frondose; attenuate flagellate branches often present 8
7. Secondary stems not stipitate or weakly so, rigidly erect; tips short attenuate or not, flagellate branches absent, microphyllous branches present or not 10
8. Secondary stem and branch leaves loosely to strongly complanate, secondary stem leaves ovate to more commonly oblong or oblong-ligulate, apex mostly acute to acute-rounded, often coarsely toothed at apex Neckeraceae p.p.
8. Secondary stem and branch leaves mostly teretely foliate; leaves ovate to ovate-short or -long acuminate 9
9. Secondary stem leaves abruptly long acuminate from a broad ovate base; margins serrulate to near base Rigodiaceae
9. Secondary stem leaves gradually short acuminate; margins entire to serrulate at apex
 Leptodontaceae p.p.
10. Leaves bordered with narrowly linear cells and conspicuously differentiated from inner laminal cells; calyptra mitrate or campanulate and fringed with ciliate hairs Daltoniaceae p.p.
10. Leaves lacking a border, marginal cells not differing greatly from inner laminal cells; calyptra mitrate or cucullate, base lacking hairs 11
11. Secondary stem leaves loosely to strongly complanate 12
11. Secondary stem leaves terete foliate 13
12. Leaves broadly ligulate, apex truncate; costa ca. 3/4 leaf length; attenuate branches uncommon; gemmae absent Neckeraceae p.p.
12. Leaves broadly elliptical, apex acute; costa percurrent; stem and branch tips short attenuate with gemmae in axil of reduced leaves Adelotheciaceae
13. Microphyllous branches present; median laminal cells fusiform to fusiform-rhomboidal; leaf margins plane; seta elongate, capsules exserted Leptodontaceae p.p
13. Microphyllous branches absent; median laminal cells oval to oblong-oval; leaf margins recurved or plane; sporophytes several to many along one side of stem, seta very short, capsules immersed Cryphaeaceae p.p.
14. Leaf costa ca. 1/2 leaf length or less; laminal cells thin-walled; plants rather delicate, stems readily breaking when removing leaves 16
14. Leaf costa mostly 2/3 leaf length or longer; laminal cells rather thick-walled; stems resilient 16
15. Leaf margins denticulate, ciliate or entire; peristome present or absent; plants common Fabroniaceae p.p.
15. Leaf margins serrulate above alar region; peristome absent; plants rare Myriniaceae p.p.
16. Upper laminal cells subquadrate; costa strongly flexuose distally Anomodontaceae p.p.
16. Upper laminal cells rhombic to short rhomboidal or oval; costa straight or weakly flexuose 17
17. Branches ascending; leaf margins entire; costa often spurred below; exostome teeth ca. 1/2 the length of the endostome Regmatodontaceae
17. Branches spreading or creeping; leaf margins dentate or serrulate; costa entire, not spurred; exostome subequal to or larger than endostome 18

18. Leaf apex relatively broad, acute; seta roughened or scabrous; capsule ovoid, to 1.1 mm long

Myriniaceae p.p.
18. Leaf apex relatively abruptly short to somewhat long acuminate; seta smooth; capsules cylindrical, mostly greater than 2 mm long Leskeaceae p.p.