MIelichhoferiaceae. Genus Pohlia

1. Plants medium-sized or small and slender, dioicous; gemmae present in leaf axils 2

— Plants large or medium-sized, rarely slender, dioicous or paroicous; gemmae absent in leaf axils 11

2. Gemmae solitary in leaf axils, opaque 3

— Gemmae numerous in leaf axils, translucent or +opaque 5

3. Gemmae spheric to elliptical; gemmae leaf primordia restricted to apices; plants slender, dull 12. P. filum

This species is widespread in Russia: Arctic, Subarctic, northern taiga regions, the Caucasus, mountain regions of South Siberia, Far East, and a few localities in central European Russia. Pohlia filum often grows on bare, sandy soil along rivers, stream banks, landslides, and roadsides. It is less commonly encountered on fine soil in rock crevices and between boulders in tundra, meadow vegetation, and willow stands. Its main diagnostic feature is the presence in the leaf axils of solitary, large, spherical, opaque gemmae with leaf primordia restricted to the apices. It can be separated from P. drummondii by the following features: gemmae spherical to elliptical vs. cylindrical to oblong; gemmae leaf primordia restricted to apices vs. scattered along the gemmae body; and plants slender, dull vs. medium-sized, glossy. Pohlia andalusica is primarily distinguished from P. filum by having gemmae numerous rather than solitary in the leaf axils and obconic form of gemmae (vs. spherical form of gemmae).

— Gemmae cylindrical to oblong; gemmae leaf primordia present at least to middle of gemmae; plants medium-sized, glossy 4

4. Plants green or dark-green; leaves ovate-lanceolate; gemmae dark-brown to black 11. P. drummondii

This Arcto-alpine species is known in Russia from northern regions, the Caucasus, Urals, mountains of South Siberia and Far East. It grows on bare soil in disturbed habitats, rock outcrops, stream/ river banks, and roadsides. It characteristically has: medium-sized, glossy plants; +strong, dark-red or black stems; solitary, large, oblong to shortly cylindrical, opaque, dark gemmae with leaf primordia at the apices and scattered along the gemmae body.

— Plants pale-whitish; leaves narrowly lanceolate; gemmae cherry-red to dark-red 13. P. beringiensis

This Arcto-alpine species is sporadically distributed in Asian Russia: Yamal Peninsula, Taimyr, the Putorana Plateau (East Siberia), Altai, Zabaykalsky Territory, Yakutia, Severnaya Zemlya Archipelago, Vrangel Island, Chukotka, and Magadan Province; it is also known from a few localities in the Arctic regions of European Russia. Pohlia beringiensis grows on bare soil slopes, fine-grained substrata in rock fields and in wet tundra. It can be recognized by its pale-whitish plants that have imbricate, narrowly lanceolate leaves, and solitary, large, oblong to shortly cylindrical, opaque, cherry-red gemmae with leaf primordia at the apices and scattered along the body. It is similar to P. drummondii, but differs in plant color (pale-whitish vs. green or dark green) and gemmae color (cherry-red vs. dark brown to blackish).

5(2). Gemmae 2–4(–7) in leaf axils, +opaque when old, with leaf-like primordia 6

— Gemmae 4–10(–15) in leaf axils, translucent when old, with peg-like primordia 8

6. Gemmae narrowly to broadly cylindrical 16. P. tundrae

In Russia this Arcto-alpine species is known from the Yamal Peninsula, Kamchatsky and Zabaykalsky Territories, the southern Far East. It grows on bare soil over upturned fallen tree roots, among exposed roots in spruce forests, and along roadsides. It is characterized by the presence of narrow or broad cylindrical gemmae with leaf-like primordia near the apices and scattered along the body. Pohlia tundrae is very similar to P. proligera in having glossy leaves and numerous narrow, elongate gemmae that give the distal parts of stems a “disheveled” appearance. They differ in several gemmae features: size (350–800 vs. 150–250 mm long); leaf primordia shape (leaf-like vs. peg-like); and leaf primordia width at base (3–6 vs. 1–3 cells).

— Gemmae obconic, oblong or spheric  7

7. Gemmae obconic to oblong; leaf primordia erect or spreading 15. P. andalusica

This boreal species occurs in central and north regions of European Russia; it is known a few localities in the Caucasus, Western Siberia, Tyva Republic and Far East. It grows on bare sandy soil in forests, on river banks, roadsides, bluffs, sand-quarries, rarely in tundra vegetation and on decayed wood in forests. Its main diagnostic features are the presence of 2–5 obconic gemmae with acute, leaf-like primordia in the leaf axils. It differs from P. bulbifera in having erect rather than incurved, dome-shaped leaf primordia on the gemmae apices.

— Gemmae spheric; leaf primordia incurved, forming a dome over gemmae apices 14. P. bulbifera

This boreal species is widespread in northern and central regions of European Russia, it is known from Siberia and Far East. It grows on bare soil usually on river/lake banks, bluffs, landslides, or roadsides. It is less frequent in tundras and forests on exposed roots or fallen trunks and occasionally found in boggy meadows/hillocks. It characteristically has slender plants and numerous (2–6), spheric gemmae with incurved leaf-like primordia in the leaf axils. Pohlia filum is the only other Russian species with spheric gemmae, but its gemmae are solitary in the leaf axils, while P. bulbifera has multiple gemmae in its leaf axils.

8(5). Gemmae linear to vermicular; plants robust, glossy  18. P. proligera

In Russia this species is distributed in the Arctic, Subarctic and boreal zones, in mountains of the Caucasus, Siberia and Far East. It typically grows on bare soil on river/lake banks, bluffs, landslides and roadsides; less commonly on fine soil in rock fields, rock crevices, and tundras. It is recognized by the presence of robust, glossy plants with dense clusters of linear to vermicular, transucent gemmae.

— Gemmae isodiametric, obconic, oblong or linear; plants large or small, dull or shiny 9

9. Gemmae on single plants various in shape (obconic, oblong or linear); plants slender, dull 17. P. annotina

Pohlia annotina is sporadically distributed in central and northern regions of European Russia, it is known from the Urals, Khanty-Mansi Autonomous Area, south Siberia and Far East. It typically grows on bare, sandy soil on stream banks, bluffs and sand-quarries, less commonly on bare soil or exposed roots in forests, and rarely on fine soil in cliff crevices. Its main diagnostic features are: slender, dull plants; gemmae on single plants variable in shape, (obconic, oblong or linear gemmae); and gemmae with 2–5 peg-like apical primordia. It is distinguished from P. proligera by: plant size (slender vs. robust); plant sheen (dull vs. glossy); and gemmae shape variation on single plants (variable, oblong to linear vs. uniform, linear to vermicular gemmae). It differs from P. andrewsii in: plant size (slender vs. robust); plant sheen (dull vs. shiny); and gemmae color (dull orange vs. bright orange, yellow or red).

— Gemmae on single plants uniform in shape (isodiametric, obconic or oblong); plants large or small, dull or shiny 10

10. Gemmae 50–120 mm long, isodiametric or nearly so, light-green to yellowish, on uni- or bicellular stalk; plants slender, dull  20. P. campthotrachela

In Russia Pohlia camptotrachela is restricted to a few localities in western and central (Leningrad, Tver, Moscow, Ryazan, and Vladimir Provinces) and northwestern (Murmansk Province and Karelia) regions of European Russia; it is known from a few localities in the Ural and Yamal-Nenets Autonomous Area. It grows on bare soil on roadsides and ditches. It can be recognized by the presence of slender, dull plants that have small, isodiametric, uniformly shaped, translucent, light-green gemmae. It is distinguished from P. annotina by several gemmae features: shape variation on single plants (uniform vs. variable); overall shape (isodiametric vs. obconical, oblong to linear); and size (50–120 vs. (100–)250–400(–600) mm long). Pohlia camptotrachela differs from P. andrewsii in having slender, dull vs. robust, shiny plants and light-green vs. orange or red gemmae.

— Gemmae 70–200 mm long, obconic or oblong to spherical, orange, yellow or red, sessile; plants robust, shiny 19. P. andrewsii

This arctic-alpine species occurs primarily in tundras and mountains of the Russian Arctic/Subarctic and montane regions of South Siberia, Yakutia and the Far East. It also occurs occasionally in north-east of European Russia (Timan Mountains of Komi Republic), the Caucasus, South Urals. Pohlia andrewsii grows in tundras on fine soil in rock fields/crevices, and bare soil in forests/meadows. It can be recognized by its robust, shiny plants that have oblong or spheric, bright orange gemmae.

11(1). Leaf margins strongly recurved nearly throughout 12

— Leaf margins plane or slightly recurved below 13

12. Stems green; leaves ovate-lanceolate; leaf cells short-rhomboidal, thick-walled; capsules erect  5. P. cardotii

Pohlia cardotii is mainly a North American species recently found on the Kamchatka Peninsula. It grows on bare soil in disturbed habitats, rock outcrops and stream banks. Pohlia cardotii is recognized by its erect capsules, strongly recurved leaf margins, and short-rhomboidal, thick-walled leaf cells.

— Stems red; leaves long-lanceolate; leaf cells linear-hexagonal, thin-walled; capsules +inclined ..4. P. crudoides

This arctic-alpine species in Russia occurs primarily in the Arctic/Subarctic regions and in mountain regions of Siberia/Far East, and in the Urals. It grows on wet rock outcrops, in tundras, and on bare soil. Its distinguishing features are: shiny green or golden plants; red stems; and narrowly lanceolate leaves with strongly recurved leaf margins.

13. Leaf cells broadly rhomboidal, mostly more than 14 mm wide 14

— Leaf cells hexagonal to linear, mostly less than 12 mm wide 17

14. Leaves obtuse, slightly cucullate; plants paroicous  6. P. obtusifolia

This arctic-alpine species in Russia is known from some localities in the Arctic/Subarctic regions of European Russia and montane regions of the Caucasus, Urals, Yakutia, Commander Islands, Chukotka. It grows on wet soil on river/stream banks, in tundras, and fine soil between boulders in rock-fields. Its main diagnostic features are: pale-green to blackish plants; crowded or distant, obtuse leaves; lax, thin-walled leaf cells; and partly reduced peristomes.

— Leaves +acute, not cucullate; plants dioicous 15

15. Plants slender; stems less than 1 cm long; setae short and thick 25. P. melanodon

Pohlia melanodon is common in central and north regions of European Russia and also present on Urals, Caucasus, in eastern Siberia (Tomsk and Tyumen Provinces). It grows on wet soil on roadsides, river banks, and in disturbed habi­tats. It is recognized by the following features: glossy, yellowish, delicate plants; lax, broadly rhomboidal leaf cells; and small, short-pyriform capsules with dark peristomes and deeply immersed stomata. It differs from P. vexans and P. brevinervis in having wider leaf cells (13–)15–27 vs. 6–13 mm wide); from P. atropurpurea it differs in deeply immersed stomata.

— Plants large; stems 1–3(–10) cm long; seta long and thin 16

16. Plants pinkish; leaf apices broadly acute to obtuse; stomata superficial 23. P.>P. ludwigii

This Arcto-alpine species is known in Russia from a few localities in Murmansk and Vologda Provinces, Novaya Zemlya, Komi Republic, Urals and Primorsky Territory. It grows on wet soil in tundra and on stream banks. Its main diagnostic features include: pink colored plants; leaves +long-decurrent; leaf apices blunt; and leaf cells thin-walled, 12–18(–25) mm wide, becoming narrower at margins forming an indistinct border. Bryum cryophyllum differs from P. ludwigii in having leaf margins distinctly bordered; Bryum weigelii differs in having considerably longer leaf decurrencies; Pohlia wahlenbergii differs in having more acute leaves with shorter leaf decurrencies and whitish or glaucous-green rather than pink colored plants.

— Plants whitish or glaucous green; leaf apices acute; stomata deeply immersed 24. P. wahlenbergii

This species is widespread and known from most regions of Russia, except the southern steppe zone. It grows on wet soil on stream/river banks, in tundra, meadows, and flood plains. It characteristically has robust, dull, whitish or glaucous-green plants and broad leaf cells (20–25 mm wide). Pohlia cruda differs from it in having plants with a distinct metallic sheen and narrower leaf cells (8–10 vs. 20–25 mm wide).

17(13). Plants with distinctive metallic sheen, pale whitish-green; stems 3–5 cm long; perichaetial and upper stem leaves long and narrow 7. P. cruda

Pohlia cruda is common and widespread throughout most of Russia, however in lowland European Russia it is sporadically distributed. It grows on bare soil in disturbed habitats, on rock outcrops, river banks, and rarely on decaying wood. The species is recognized by its pale whitish or glaucous-green plants that have a strong, distinct metallic sheen; elliptic leaves with plane margins and long, thin-walled cells; cylindric-pyriform capsules; perfect, well-developed endostomes with long cilia.

— Plants without metallic shine; other character state combination not exactly as above 18

18. Rhizoidal tubers present 19

— Rhizoidal tubers lacking 20

19. Rhizoidal tubers orange to brown, round to ovoid, superficial cells plane 21. P. lescuriana

This species is sporadically distributed in northern/central European Russia and East Siberia; it is known from a few localities in east Siberia and Chukotka. It grows on wet, bare soil in disturbed habitats, on roadsides and river banks. Pohlia lescuriana is characterized by its: delicate, often whitish plants; short, broadly pyriform capsules; superficial stomata; yellow exostome teeth; and orange to brown rhizoidal tubers. Pohlia vexans and P. melanodon differ from P. lescuriana in having immersed rather than superficial stomata; P. atropurpurea differs from it in having dark brown rather than yellow exostome teeth.

— Rhizoidal tubers pale-yellow, stellate, superficial cells bulging 22. P. lutescens

Pohlia lutescens has only recently been found in Russia but it has a widely scattered distribution in the Urals (Perm) and in the Far East (Amur Province and Khabarovsk Territory). It grows in conifer forests on bare soil over upturnd roots of fallen trees, and river banks. Pohlia lutescens can be recognized by its very delicate plants and pale-yellow, stellate rhizoidal tubers. Pohlia lescuriana differs from it in having round to ovoid rather than stellate rhizoidal tubers that are orange to brown rather than yellow in color.

20. Plants whitish; leaf margins recurved; leaf cells linear, very thin walled; capsules pale 10. P. alba

Pohlia alba is known only from its 19th century type collections made on the lower course of the Yenissey River (northeastern Siberia). Its distinctive features include: whitish plants; light-colored, ovate, pendent capsules; yellow exostome teeth; and endostomes with perforated segments and long cilia.

— Plants with character state combination not exactly as above 21

21. Capsules short-pyriform to hemispherical; exothecial cells +isodiametric with sinuose walls; plants dioicous 22

— Capsules elongate-pyriform to narrowly cylindrical; exothecial cells rectangular with straight walls; plants paroicous or rarely dioicous (P. sphag­nicola, P. saprophila) 24

22. Exostome teeth yellow; endostome hyaline; stomata strongly covered by subsidiary cells 27. P. vexans

Pohlia vexans is known from a few localities in Russia (Arkhangelsk Province, Nenets Autonomous Area, the Altai, Chukotka, Khabarovsk and Primorsky Territories). It grows on wet, bare soil on river banks and in disturbed habitats. It is characterized by its: delicate, glossy plants; red stems; erect leaves; linear-rhomboidal, thin-walled leaf cells; short, pyriforme capsules; and pale exostome teeth. Pohlia vexans differs from P. atropurpurea and P. brevinervis in having pale vs. brown exostome teeth and lighter colored endostomes; and stomata strongly vs. not or slightly covered by subsidiary cells. Pohlia melanodon differs from P. vexans in have dark-brown rather than pale exostome teeth and linear-rhomboidal rather than widely rhomboidal leaf cells. Pohlia lescuriana differs from P. vexans in having mostly superficial rather than immersed stomata and rhizoidal tubers (lacking in P. vexans).

— Exostome teeth dark-brown or red-brown; endostome yellow; stomata superficial or slightly covered by subsidiary cells 23

23. Plants dark-green to reddish; leaf cells rhomboidal, 10–15 mm wide 26. P.>P. atropurpurea

This northern species is known in Russia from north regions of European Russia, Yamal and Taimyr Peninsulas, Tyumen and Magadan Provinces. It grows on wet, bare soil on river banks and roadsides. It is characterized by delicate, dark, often reddish plants; shortly pyriform to hemispherical capsules; and dark-brown exostome teeth.

— Plants whitish; leaf cells linear-rhomboidal, 6–9 mm wide 28. P. brevinervis

This very rare species is known only from a few localities in North America and the Yenissey River in northern Siberia. It is distinctive in having whitish plants; shortly pyriform capsules; and dark-brown exostome teeth. Pohlia brevi­nervis is very similar to P. atropurpurea; they dif­fer in cell width (6–9 vs. 10–15 mm wide) and plant color (whitish vs. dark-green or reddish).

24(21). Endostomial cilia long, nodose 25

— Endostomial cilia short, nodes not thickened or cilia absent 27

25. Plants paroicous 8. P. nutans

Pohlia nutans is widespread throughout Russia, and very common in the forest/tundra zones. It grows on soil, decaying wood, and tree bases, in tundras, on mire hummocks, rock outcrops, and bare soil in disturbed habitats. The species can be recognized by several diagnostic features: dark-green plants; thick-walled leaf cells; paroicous sexual condition; numerous long-pyriform, pendent capsules; and long, nodose endostomial cilia.Pohlia nutans is most similar to P. elongata. They differ primarily in sporophytic features: capsule shape (long- vs. short-pyriform) and endostomial cilia length (long vs. short or absent). Unfortunately, when sporophytes are absent it is very hard to separate them.

— Plants dioicous 26

26. Leaf margins weakly serrulate to entire, plane; capsules pyriform 9. P. sphagnicola

The distribution of this species is difficult to assess because it is often confused with P. nutans. Pohlia sphagnicola differs from P. nutans in sexual condition (dioicous vs. paroicous) and in having weakly serrulate to entire rather than clearly serrulate upper leaf margins. As implied by its name P. sphagnicola grows only among Sphagnum in mires, but P. nutans is occasionally found in this type of habitat.

— Leaf margins denticulate, recurved; capsules narrowly cylindrical 3. P. saprophila

This rare Asian species is known in Russia from the Altai Mountains, Zabaikalsky Territory, Republic of Buryatia, and Kamchatka. It grows on decaying wood in forests. Pohlia saprophila can be recognized by its growth in compact tufts; dense, tomentose rhizoids; light-green plant color; long-lanceolate leaves with narrowly recurved margins; narrowly cylindrical capsules; and dioicous sexual condition.Pohlia longicollis and P. elongata differ from P. sapro­phila in: growth form (loose vs. compact tufts); sexual condition (paroicous vs. dioicous); and endostomial cilia length (short or absent vs. long). Pohlia sphagnicola differs from P. saprophila in growth form (separate, individual plants vs. compact tufts) and capsule shape (pyriform vs. narrowly cylindrical).

27. Plants dark-green, dull; leaves broadly lanceolate; leaf cells 50–85 mm long, rhomboidal, thick-walled 1. P. elongata

Pohlia elongata is common in the mountains of Siberia, Far East, and the Caucasus; it is also known from the Urals and Komi Republic. The species grows on rock outcrops and bare soil in disturbed habitats. Its main distinguishing features are very long capsules; thick-walled leaf cells; and short stems. Pohlia longicollis differsfrom P.elongata in having longer leaf cells (50–150 vs. 35–85 mm) with thin rather than thick cell walls; and silky, bright green rather than dull, dark-green plants. Pohlia nutans differs from P. elongata in having pendent rather than inclined capsules with shorter necks, and long rather than short or absent endostomial cilia. The distinc­tions between P. elongata and P. saprophila are discussed under the latter species.

— Plants green, shiny; leaves long-lanceolate; leaf cells 80–140 mm long, linear-rhomboidal, thin-walled 2. P. longicollis

Pohlia longicollis is a common, mountainous species known in Russia from Siberia and the Far East, Murmansk Province, Komi Republic, the Caucasus, and Urals. It grows on rock outcrops and bare soil in disturbed habitats. The species can be recognized by its shiny, pure-green plants; crowded, long-lanceolate leaves; long, linear-rhomboidal, thin-walled leaf cells; long capsules. It differs from P. crudoides in having plane leaf margins, and from P. cruda by its paroicous sexual condition, lack of a strong metallic plant sheen, and longer, crowded leaves. The distinctions between P. longicollis and P. sapropila are discussed under the latter species.