Orthotrichiaceae. Genus Ulota

1.Mature capsules pyriform, shortly 8-ribbed at mouth, smooth below; capsule mouths irregularly and strongly puckered2. U. coarctata

This species is quite distinct in having, nearly smooth, pyriform capsules that are strongly puckered at the mouth. Remarkably, this odd feature occurs convergently in Orthotrichum and Macromitrium. Additional important features of U. coarctata include: long-acuminate, curved leaves; short, non-fused exostome teeth; and mostly absent endostomial segments. In RussiaU. coarctata occurs in Kaliningrad Province and at moderate elevations (530 to 1250 m) in the Western Caucasus. It is a corticolous moss that grows on beech and alder.

Mature capsules oblong-cylindrical, fusiform, urceolate or obconic, 8-ribbed throughout; capsule mouths evenly contracted 2

2.Dry capsules fusiform, narrowed toward the mouth 3

Dry capsules oblong, cylindrical, urceolate or obconic, often strongly contracted below mouth 5

3.Leaves flexuose to loosely erect when dry; endostome absent6. U. drummondii

This species can be distinguished in the field by the following combination of features: curved to loosely erect leaves; fusiform capsules gradually narrowed to the mouth and strongly furrowed above resulting in irregularly angular to star-like capsule mouths; and single, whitish peristomes with often fragile and broken exostome teeth directed upward to spreading. In Russia U. drummondii is mainly a Far Eastern species found in the humid, oceanic areas of Sakhalin and Kuril Islands, as well as western and southern Kamchatka. In some areas, e.g. Iturup Island (Kuril Islands), it is the most common species of the family. There is a single collection of U. drummondii from one of the Baltic islands in European Russia. Ulota drummondii mostly grows on willow, poplar and alder, often with other Ulota species and/or Lewinskya sordida.

Leaves crisped-contorted when dry; endostome present 4

4.Endostome segments mostly smooth; spores 2732 mm; Asian species [U. delicata]

Ulota delicata was described from China (Fujian Province), and is also known from Japan (Honshu). Although it has not yet been found in Russia the species could occur in the southern Russian Far East. Indeed, some collections from the coastal areas of Primorsky Territory and Kamchatka have similar contorted to crisped leaves and fusiform capsules. However, these collections have either young sporophytes or old, eperistomate capsules and differ from U. delicata in having acuminate rather than obtuse to acute leaves and smaller spores.

Endostome segments distinctly longitudinally striolate; spores 2024 mm; European species 1. U. bruchii

The presence of fusiform capsules that are narrowed to the mouth, distinguish U. bruchii from U. intermedia and U. crispula.Ulota bruchii differs from U. crispa s. str. in having irregularly reflexed exostome teeth that are often split in mature capsules. Garilleti et al. (2000) indicate the best way to separate U. bruchii from U. crispa s. lat. is the presence of longitudinally striolate endostome segments.Ulota bruchii is a European Russia species found in Kaliningrad and Leningrad (Baltic Sea islands) Provinces.

5(2). Basal juxtacostal cells rectangular, thin- to moderately thick-walled, longitudinal walls straight, non- or weakly sinuose, non-porose; leaves mostly flexuose to loosely erect 6

Basal juxtacostal cells elongate, strongly thick-walled, longitudinal walls more or less sinuose, often porose; leaves straight, flexuose to loosely erect, or crisped 10

6.Perichaetial leaves differentiated; perichaetial leaf apices obtuse to rounded 7

Perichaetial leaves not differentiated; perichaetial leaf apices acute to acuminate 8

7.Capsules obconic to cylindric, rarer oblong, not contracted below mouth, brown; exostome teeth 16, 380510 mm long 10. U. orientalis

Ulota orientalis was recently described from the Khabarovsk Territory (Sovetskaya Gavan); it grows on alder trunks on steep, wind-blown, sea-facing slopes (Fedosov & Ignatova, 2018). It differs from other Holoarctic Ulota species (except U. japonica, U. pacifica and U. barclayi Mitt.) in having small spores and somewhat thin-walled, rectangular, basal juxtacostal leaf cells with longitudinal walls that are not sinuose or porose. It is similar to U. megalospora Venturi and U. rehmannii in having obconic capsules that are gradually narrowed from the mouth to the setae and striolate on the upper, inner surface of the exostome teeth. Ulota orientalis differs from U. japonica in having perichaetial leaves obtuse to rounded (vs. acute); capsules brown, oblong to obconic not constricted below the mouth (vs. yellowish to light brownish, mostly urceolate, markedly constricted below the mouth); bands of exothecial cells weakly differentiated (vs. strongly differentiated); exostome teeth much longer, 16 (vs. shorter, mostly 8); and endostome segments slender, non-keeled, composed of one cell row (vs. robust, keeled, composed of two cell rows). For the distinctions between U. orientalis and U. pacifica see discussion under the latter species.

Capsules urceolate, cylindrical to ovate, mostly contracted below mouths, yellow to brownish; exostome teeth in 8 pairs, up to 350 mm long [U. barclayi]

Vitt (2014) considered this species closely related to U. japonica. Indeed, the two species were treated as conspecific (Anderson et al., 1990); however Garilleti et al. (2015) and Caparros (2015) accepted it as a good species. It is a western North American species described from Sitka Island, Alaska. Ulota barclayi is similar to U. orientalis in having differentiated perichaetial leaves, but differs from U. japonica, U. orientalis andU. pacifica in having lightly colored capsules that are contracted below the mouth; 8 pairs of exostome teeth; and red rimmed, angled opercula. The operculum characters are very useful in distinguishing U. barclayi from U. japonica when the perichaetial leaves are poorly differentiated. According to Caparros (2015) the species may occur in Asia.

8.Capsules cylindrical to ovate, slightly inflated, not contracted below mouth, weakly longitudinally ribbed; segments filiform11. U. pacifica

This recently described species from the South Kurils (Iturup and Shikotan Islands) grows on trees as well as coastal cliffs (Fedosov & Ignatova, 2018). Ulota pacifica is most closely related to U. japonica, but differs in having capsules cylindrical to ovate, slightly inflated, not contracted below the mouth, and weakly ribbed (vs. mostly urceolate, non-inflated, contracted below the mouth, and strongly ribbed); endostome segments filiform, widened only in basal­most part (vs. linear, gradually widened below); and leaf cells smooth or with a few scattered papillae (vs. regularly papillose). Ulota orientalis differs from U. pacifica in having remarkably differentiated perichaetial leaves and typically turbinate capsules. Ulota pacifica and U. curvifolia are saxicolous mosses that have somewhat short, often curved setae. But, U. pacifica differs from U. curvifolia in the following features: lower leaf cell thickness (thin-walled vs. thick-walled); upper leaf cell papillae form (absent or sparse, single vs. regularly present, mostly forked); mature capsules shape (never urceolate vs. more or less urceolate); exothecial cell band differentiation (weakly vs. strong­ly); and exostome teeth ornamentation (papillose vs. distally ridged). The East Asian U. perbreviseta Dixon & Sakurai resembles U. pacifica in having short, curved setae; capsules not constricted below the mouth; and filiform endostome segments. It differs in having crisped leaves and thick-walled lower leaf cells.

Capsules urceolate, not inflated, contracted below mouths, strongly longitudinally ribbed; segments linear 9

9.Perichaetial leaves differentiated; opercula red rimmed at base, angled when viewed from above [U. barclayi]

Perichaetial leaves not differentiated; opercula not rimmed at base, rounded when viewed from above 9. U. japonica

Wang & Jia (2012) considered U. japonica distinctive in having somewhat small plants and rectangular, more or less thin-walled basal juxtacostal leaf cells with non-sinuose, non-porose longitudinal walls. Ulota barclayi has all of these features, but differs from U. japonica in the following: perichaetial leaves (obtuse to rounded vs. acute); capsules (oblong to obconic, brown vs. mostly urceolate, light brown); exostome teeth number (16, not fused vs. 8 fused pairs). Some populations of U. japonica have weakly differentiated basal leaf cells and these can be confused with U. reptans which is also similar to U. japonica in having somewhat small plants, slightly curved leaves, papillose exostome teeth and sparsely hairy calyptra, but its basal leaf cells are clearly thick-walled. Other distinctive differences between U. japonica and U. reptans include: leaf size (1.62.7 vs. 1.01.5 mm long); capsule mouths when dry (contracted vs. mostly not contracted; and spore size (1624 vs. 2734 mm). Ulota japonica is an amphipacific, oceanic, temperate and boreal species; in Russia it is restricted to the Russian Far East: South Kurils, Sakhalin, Primorsky/Khabarovsk Territories and Kamchatka. The species is fairly common in the coastal areas of Primorsky Territory and the South Kuril Islands. It grows on the trunks and branches of willow, poplar, alder, spruce and fir in the forest belt, usually at low elevations, but occasionally as high as the timber-line.

10(5). Plants saxicolous 11

Plants corticolous 12

11.Leaves stout, mostly straight when dry; leaf cell papillae low, simple; exostome teeth on dorsal (outer) surfaces papillose throughout 7. U. hutchinsiae

This species is distinguished from all other Russian Ulota species in having saxicolous plants and leaves that are mostly straight when dry. It differs from all Russian saxicolous species of Orthotrichum in having superficial stomata. In addition its occurrence on acidic rather than calcareous rocks distinguishes it from the saxicolous O. anomalum and O. urnigerum. The genus Lewinskya is similar to U. hutchinsiae in having superficial stomata, but most Lewinskya species are corticolous or have smooth to weakly furrowed capsules with reduced endostomes (e.g., L. laevigata and L. pylaisii). In contrast, U. hutchinsiae has distinctly furrowed capsules with well-developed endostomes. In Russia U. hutchinsiae is found in the western part of country: southern Karelia Republic, Berezovij Island in the Baltic Sea, Leningrad Province, and the Caucasus (Krasnodar Territory and North Ossetiya). The species is also known from a highly disjunct locality along the middle course of the Biya River in the Altai Mts (Bardunov, 1974; Ignatov & Ochyra, 1994).

Leaves slender, contorted when dry; leaf cell papillae high, partially forked; exostome teeth on upper dorsal (outer) surfaces longitudinally ridged 5. U. curvifolia

Ulota curvifolia can be distinguished from all other Holarctic Ulota species by the following combination of features: plants saxicolous; leaves contorted when dry; leaf cell papillae high, partially branched; capsules subspherical to short-urceolate; calyptrae densely hairy; and exostome teeth longitudinally striolate on upper dorsal (outer) surfaces. In addition, U. curvifolia has more a northern distribution and/or occurs in higher elevations than all other Ulota species. It is a widespread species in Russia and the only Ulota species found in the subarctic and continental mountain areas wherever acidic crystalline bedrocks occur. On the other hand it is unknown in boreal mountain regions composed of basic to neutral rocks (Putorana Plateau) and volcanic areas (Kamchatka). It is especially abundant on shaded overhanging granitic and gneiss rocks where it usually forms extensive mats; occasionally it occurs on meta-gabbro.

12.Leaves strongly contorted or crisped when dry; calyptrae densely hairy 13

Leaves flexuose to contorted, calyptrae naked or sparsely hairy 17

13.Endostome segments striolate below1. U. bruchii

Endostome segments mostly smooth below 14

14.Leaves contorted to slightly crisped when dry; leaf bases scarcely widened, weakly concave, elliptic, gradually narrowed to acumina; basal marginal leaf cells weakly differentiated in 13 rows; endostome segments uniseriate or with biseriate patches, transverse cell walls thin4. U. crispula

This species differs from U. crispa and U. intermedia in having contorted to weakly crispate leaves; a weakly differentiated, narrow, group of cells along the basal leaf margins; a different exothecial cell band structure; and partly biseriate endostome segments (see Caparros et al., 2016). Ulota crispula can be confused with Ulota species not related to U. crispa because of the presence of leaves that are weakly widened at base and contorted rather than crisped. It differs from U. japonica in thick-walled basal leaf cells and mostly larger spores. It differs from U. barclayi, U. rehmanniiandU. reptans in having capsules that are contracted below the mouth and weakly differentiated basal marginal leaf cells. In Asian Russia some specimens are hard to place in either U. intermedia or U. crispula because they are gametophytically similar to U. intermedia (strongly crispate leaves; broad leaf bases; well-differentiated, broad band of basal marginal leaf cells) but sporophytically agree better with U. crispula (exothecial cell bands yellowish throughout; endostome segments fragile, partly biseriate); in addition, capsules in these plants are notably contracted below the mouth. These difficult to place, intermediate collections are widespread on the Kamchatka Peninsula and occur sporadically in the Altai Mountains, Khabarovsk Territory (Botchi State Reserve), and Far Eastern Pacific Islands (Sakhalin and Southern Kurils). In Russia U. crispula s. str. apparently occurs only in the Caucasus. The status of these intermediate Asian plants, as well as their relationship to U. crispula and U. intermedia needs to be clarified in the course of an overall revision of the U. crispa complex in the Pacific region.

Leaves strongly crisped when dry; leaf bases wide, concave, round-ovate, abruptly narrowed to acumina; basal marginal leaf cells strongly differentiated in (2)39 rows; endostome segments uniseriate, transverse cell walls thick 15

15.Mature capsules very narrow, urceolate, strongly constricted below the mouth when dry and empty; ribs separated by narrow furrows, collapsed at the constricted area of the urn; exothecial cell bands orange, 46 cell rows wi 3. U. crispa

Caparros et al. (2016) distinguish this species from the closely related U. crispula and U. intermedia by the presence of urceolate capsules that are strongly contracted below the mouth; wide exothecial cell bands with narrow furrows in between that collapse when the capsules constrict; and 8-paired exostome teeth that never split into 16 teeth. From other Ulota species it differs in having wide, concave leaf bases that abruptly taper above; and strongly crisped leaves. Caparros et al. (2016) consider U. crispa a temperate, amphioceanic species. It is widespread in southern and Middle Europe, extending northward to southern Norway and eastward to Turkey, the Caucasus and Transcarpathia. In East Asia the species occurs in China, Japan, and Taiwan. In North America it occurs along the Pacific coast (British Columbia and Washington) with a few inland localities, but seems to be absent in Atlantic North America. In Russia the species occurs in the western part of the Russian Caucasus where it is especially abundant in the southernmost Black Sea coastal area, eastward to Kabardino-Balka­ria; there are a few specimens known from Primorsky Territory in the Russian Far East. All specimens from middle European Russia, the Altai and other parts of the Russian Far East previously identified as U crispa are now referred to U. intermedia or U. cf. crispula.

Mature capsules wide, cylindrical to urceolate, not or weakly constricted below mouths when dry and empty; ribs separated by more or less broad furrows in the upper half of urn; exothecial cell bands yellowish, 25 cell rows wid 16

16.Capsule mouths reddish rimmed; exostome teeth mainly reddish with hyaline border; endostome segments 16, filiform; spores 1520(23) m[U. longifolia]

Ulota longifolia Dixon & Sakurai is a Japanese endemic usually considered a subspecies of U. crispa. It differs from U. crispa s. str. in having capsule shape and coloration similar to U. intermedia (cylindrical, not or slightly contracted below mouth, with light ribs, composed of elongate cells);longer peristome teeth (325360 vs. 320 mm long), and 16, filiform vs. mostly 8 (occasionally 16), linear endostome segments. The relationships of U. longifolia are difficult to evaluate because it exhibits a combination of features found in U. intermedia and U. crispa.

Capsule mouths not rimmed; exostome teeth pale throughout; endostome segments mostly 8, broad­ly linear; spores 1835 mU. intermedia

Ulota intermedia differs from all other Russian Ulota species in having wide, rounded to obovate, notably concave leaf bases that are abruptly narrowed to linear-lanceolate acumina; leaves strongly crispate when dry; and obconic or short-cylindrical capsules that are not contracted below the mouth. Other distinctive features of the species include: wide basal leaf borders of short rectangular cells; exothecial cell bands narrow, yellowish, with incrassate transverse walls and hyaline lumina; and endostome cells with thickened transverse walls (cf. Caparros et al., 2016). Ulota intermedia is an amphioceanic species, but its exact distribution remains unknown because for a long time it was considered conspecific with U. crispa. Ulota intermedia and U. curvifolia are the most widespread Ulota species in Russia. In fact, all specimens of the U. crispa complex in Middle European Russia are U. intermedia;eastwardit extendsto Ryazan, Vladimir and Kostroma Provinces. It is also present around Teletskoe Lake (Altai Mts.) in places with mild, wet climatic conditions. Most specimens previously named U. crispa from coastal areas of the Russian Far East are also U. intermedia. It is a corticolous species that grows on a wide array of trees: e.g., Chosenia, Populus, Salix, Betula, Picea, Abies, Acer, and Pinus pumila.

17(12). Leaves (1.4)1.82.5(2.8) mm long; upper/median leaf cells weakly papillose to nearly smooth; basal marginal leaf cells differentiated in 37 rows; exostome teeth weakly papillose or smooth, always striolate on upper, inner (ventral) surfaces 12. U. rehmannii

Ulota rehmannii can be distinguished from most other Russian Ulota species by the combination of curved leaves; strongly incrassate basal leaf cells; conic, nearly smooth calyptrae; mature, old capsules obconic; exostome teeth mostly fused in 8 pairs and striolate on upper, inner (ventral) surfaces; and large (2139 mm) spores. It is similar to the East Asian U. reptans and western North American U. megalospora Venturi in having weakly curved leaves; cylindrical to obconic capsules; large spores; and conic, nearly naked calyptrae. Ulota rehmannii differs from U. megalospora in leaf apex form (acute to short-acuminate vs. long-acuminate), stomata position (basal vs. median) and spore size (2139 vs. 3560 mm). In Russia the species is fairly common in humid, hemiboreal mountain forests from southern Siberia (Altai Mts) to the southern Russian Far East (Sakhalin and Kuril Islands).

Leaves 1.01.5(2.0) mm long; upper/median leaf cells strongly papillose; basal marginal leaf cells differentiated in 13 rows; exostome teeth papillose on upper, inner (ventral) surfaces 13. U. reptans

Ulota reptans is similar to U. rehmannii and U. megalospora in having weakly curved leaves; cylindrical to obconic capsules; large spores; and conic, nearly naked calyptrae. Wang & Jia (2012) distinguished U. rehmanniifrom U. reptans on the basis of its leaf size (1.52.8 vs. 1.01.5(2.0) mm long); and exostome teeth ornamentation (striolate vs. papillose). Ulota reptans is known from four Russian Far Eastern (Khabarovsk Territory and Sakhalin Island) collections. In addition, there are several anomalous specimens from Khabarovsk Territory that exhibit the critical features of U. reptans in transitional combinations. These collections require additional study before they can be confidently placed. Ulota reptans is a corticolous species found in hemiboreal forests on spruce, fir and alder.